Annals of the Missouri Botanical Ga ye umber Annals of the Missouri Botanical Garden Volume 91, Number 1 April 2004 The Annals, published quarterly, contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden, St. Louis. Papers originating out- side the Garden will also be accepted. All manuscripts are peer-reviewed by quali- fied, independent reviewers. Authors should write the Managing Editor for informa- tion concerning arrangements for publishing in the ANNALS. Instructions to Authors are printed in the back of the last issue of each volume and are also available online at www.mbgpress.org. Editorial Committee Victoria C. Hollowell itor, Missouri Botanical Garden Amy Scheuler McPherson Managing Editor, — Missouri Botanical Garden Diana Gunter Associate Editor, Missouri Botanical Garden Kevin Brown MBG Press Assistant, Missouri Botanical Garden Barbara Mack Administrative Assistant Ihsan A. 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The journal Novon is included in the sub- scription price of the ANNALS, annals@mobot.org (editorial queries) http://www mbgpress org The Annals are abstracted and/or indexed in A ingenta, ISI® databases, JSTOR, Research Alert © Missouri Botanical Garden 2004 THE ANNALS OF THE MissOURI BOTANICAL GARDEN (ISSN 0026-6493) is published quar- terly by the Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, MO 63110. Periodicals postage paid at St. Louis, MO and additional mailing offices. POSTMASTER: Send address changes to ANNALS OF THE MISSOURI BOTANICAL GARDEN, % Allen Marketing & Management, Р.О. Box 1897, Lawrence, KS 66044-8897. GRICOLA (through 1994), APT Online, BIOSIS®, ®, and Sci Search®, The mission of the Missouri Botanical Garden is to discover and share knowledge about plants and their environment, in order to preserve and enrich life. This paper д MN еа i 48 ( ce of Paper). Volume 91 Annals Number 1 of the 2004 Missouri Botanical Garden A REVISION OF TRISETUM. Victor L. Finot,” Paul M. Peterson,“ PEYRITSCHIA, AND 1 Soreng,? and Fernando О. SPHENOPHOLIS (POACEAE: POOIDEAE: AVENINAE) IN MEXICO AND CENTRAL AMERICA ABSTRACT A taxonomic treatment of Trisetum, Peyritschia, and Sphe 8 for Mexico and Central Americ a is given. In Mexico and Central America four species of Peyritschia. two species of | ie ded and 17 species of Trisetum s. str. are recognized. Peyritschia deyeuxioides and F P. pringlei range E Mexico to Ecuador, P. koelerioides i is found in southern lexico to Guate зне and P. humilis is mille mic to Mexico. a. е ranges from Canada to es U.S.A. and Mexico. whereas S. interrupta is found in the southwester nd Baja California, Mexico. Mexico has the ecies of Trisetum të - found in — largest e of Fl species al 15. and nine of these are Five sp E three in Costa Rica and Panama. and a single species is found i in 1 Honduras and the Dominican Republic. ew subgenus Deschampsioidea (Louis-Marie) Finot in Trisetum is propi oed. Four new species of Trisetum from Mexic o are described and illustrated: J. durangense Finot & P. M. Peterson, T. martha- -gonzaleziae P. M. Peterson & Finot, and J. Ja Hlenbergii Soreng, Finot & P. M. j erson (all in subg. Deschampsioidea ); and T. ба ашм Finot & Zuloaga (in subg. Trisetum, sect. dar pies Kevs for the genera, subgenera, sections, an id species of Trisetum, Peyr- itschia, and Си that occur in Mexico а n "entral America are given. The names Trisetum AU E Fourn. and Trisetum subsect. Desc 5 Louis-Marie are lectotypified. Key words: Aveninae, Gramineae, Peyritschia, Poaceae, Pooideae, Sphenopholis, Trisetum. This paper is part of the doctoral thesis of VLF in the Dpto. Botánica. Universidad de Concepción, Concepción. Chile. We thank Alice Tangerini and 7 Dudás E preparing E illustrations. Thanks to the Directors and ‘urators of the following herbaria: ВАА. BAF. C. CHDIR, CONC, CR, QCA, P. US, S, and SI. We thank Olof Ryding (Botanical Museum Copenhage n) for sending a xerox of и. type of Deyeuxia gracilis, and Victoria C. Hollowe ll and an anonymous reviewer for suggesting improvements to the manuse ript. VLF gratefully ac ‘knowledges the Pedersen Foundation for a fe llowship to КИР types of Trisetum at Paris (Р) апа Stoc 'kholm (S); the MINEDUC for a fellowship to study types at the Smithsonian Institution. Washington. D.C.. and the Instituto de Botánica deque cs nos Aires; the financial suka from Dirección de Investigacion, Universidad de Concepción, Project DIUC 201.121.005- L. 0; and the help of Oscar Matthei and Clodomiro Marticorena (U. de Concepción, Chile) for directing his Ph.D. 2 Unive rsidad de Concepción. Facultad de Agronomía. Dpto. Producción Animal. Casilla 537, Chillan. Chile. vifinot@udec.cl. ir epartment of Botany. National Museum of Natural History, Smithsonian Institution, Washington. D.C. 20013-7012, ra de Botánica Darwinion, Labardén 200, San Isidro 1642, Argentina. ANN. MISSOURI Bor. GARD. 91: 1-30. 2004. Annals of the Missouri Botanical Garden RESUMEN Se presenta un tratamiento taxonómic o de los géneros Trisetum, Peyritschia у Sphenopholis para шек у América Central. Cuatro especies de sch nocidas en México y Ce Шш гїса. Peyritse hia deye >UXLOL rioides se encuentra uen el sur ientra en Canadá, EE. we se епс! de Trisetum se encuentran en Guatemala. tres en Ce Se propone un nuevo subgénero Desc hampsioide a " ш Меле ») Finot en Trisetum. Cuatro nue dos especies de Sphenopholis у йез ур, México, mientras 5. in terrupta habita el sudoeste de EE. México. México posee el mayor nümero de especies Е Trisetum, 15 especies, nueve de ellas endémicas. Ci a Rica y Panamá y sólo una en Honduras y Re públic a Hone 'ana. . fueron reco- E koe 17 especies de Trisetum s. pringlei habitan desde México x e aor Ub. у n 1 мис Inco espec ies de México son descritas e ilustradas: T. durangense Finot & P. M. Peterson. T. martha-gonzaleziae P. M. Peterson & Finot y T. spellenbergii Sore ng. Finot & P. M. Peterson (todas en subg. Desc "hampsioidea); v T. ligulatum m & Zuloaga (en subg. Trisetum, sect. Trisetaera). Se presentan c ru para los géneros, subgéneros, secciones y especies de Trisetum, Peyritschia y cox bana nr presentes en México у América Central. Los elie Trisetum gracile Е. ЫШ. у Trisetum subsect. Desc hampsioidea Louis-Marie fueron lec totipificados. Trisetum Pers. comprises about 70 species in temperate regions of Europe, Asia, America, Aus- tralia, and New Zealand (Louis-Marie, 1928-1929: Hitchcock, 1939; Swallen, 1948; Tsvelev, 1970. 1983: Jonsell, 1980; Veldkamp & Van der Have, 1983: Clayton & Renvoize, 1986; Pohl & Davidse, 1994, 2002; Edgar, 1998; Soreng et al., 2003). A general account of the American taxa of Trisetum was published by Louis-Marie (1928-1929). in which he recognized a total of 60 species. Hitch- cock (1939) recognized 19 species of Trisetum in North There are 38 species of Trisetum in North. Central. and South America (Finot, 2003b). ception of the cosmopolitan J. spicatum (L.) K. Richt., stricted geographical distribution. The majority of America, including Mexico. With the noteworthy ex- most species of Trisetum have a rather re- — the species are from Mexico, where 9 are endemic, and only J. irazuense (Kuntze) Hitche. extends from Mexico and Central America to Ecuador in South America. The systematic relationships of the Mexican and Central American species of Trisetum are not well known, in part due to the close affinity with Pey- rischia E. Fourn. and Sphenopholis Scribn. Four- nier (1886) one species from Mexico and Guatemala, P koele- Fourn. Hitchcock (1939) also rec- his treatment of the North described the genus Peyritschia with rioides (Peyr.) E. ognized Peyritschia American flora. Koch (1979) transferred to Peyrits- chia a second species from Mexico, Costa Rica, and Seribn. [= Р. Hernández-Torres Guatemala, Deschampsia pringlei pringlei (Scribn.) S. D. and Koch (1987) did not recognize Peyritschia and Koch]. placed both of these species in Trisetum. The sys- tematic position of these two species Trisetum has not gained universal acceptance, although some recent treatments recognize Peyritschia (Clayton & Renvoize, 1986: Watson & Dallwitz, 1992; Finot. 2003a). One further species, Peyritschia conferta Pilg.) Finot, occurs in Venezuela, Ecuador, and Bo- — livia. Peyritschia differs from Trisetum in having 1- nerved glumes, lemmas with bilobed apices that are awnless, mucronate, or with a well-developed awn borne near the base, paleas that are tightly en- closed by the margins of the lemma, and florets with only two stamens. Trisetum has a first glume l- to 3-nerved, a second glume 3- to 5-nerved, lem- ma apices with 2 to 4 short awns with the central awn inserted above the middle of the lemma, paleas that are not tightly enclosed by the lemma (i.e.. gaping), and florets with 3 stamens. Sphenopholis was described by Scribner (1906) to include seven species and seven subspecies. He characterized the genus as having spikelets that disarticulate below the glumes and between the flo- rets, second glumes that are usually broadly ob- ovate, and lemmas with an entire or 2-toothed apex that is awnless or awned just below the apex. In contrast, Trisetum has spikelets that disarticulate above the glumes and between the florets, second glumes that are lanceolate to ovate, and lemmas with a biaristulate to bidentate apex that is awned on the upper third. On the basis of these characters, Scribner (1906) transferred three species of Trise- tum to the new genus Sphenopholis: S. interrupta (Buckley) Seribn.. S. hallii (Scribn.) Seribn.. and S. palustris. (Michx.) Scribn. Hitchcock (1939) and Hitchcock and Chase (1951) retained these three species in Trisetum, although Hitchcock (1915) ear- lier placed S. palustris as a synonym of a fourth species, S. pensylvanica (L) Hitche. Erdman (1965) also recognized S. pensylvanica but did not include 5. interrupta, although he recognized that this spe- cies resembles Sphenopholis in spikelet disarticu- lation. Volume 91, Number 1 Finot et al. 3 8 5 Peyritschia & Sphenopholis Steudel (1854). made one of the earliest enumerations of Trisetum who recognized four species. "p in Mexico: T. deyeuxioides (Kunth) Kunth, T. elon- gatum (Kunth) Kunth, T. tolucense (Kunth) Kunth, and T. viride (Kunth) Kunth. An important contri- bution to the knowledge of Mexican species was made by Fournier (1886). who described six new species: T. gracile E. Fourn., T. nivosum E. Fourn., T. interruptum E. Fourn., T. paniculatum E. Fourn.. T. bambusiforme E. Fourn.. and T. virletii E. Fourn. On the basis of Pringle's collections from Mexico. (1896) Grasses of North America T. deyeuxioides var. pu- Scribner in Beal's described three new taxa: bescens Scribn. ex Beal, T. filifolium Scribn. ex Beal var. filifolium, and T. filifolium var. Scribn. ex Beal. In a general synopsis of Mexican Hitchcock (1913) described J. palmeri Hitche. and mentioned nine other species. By the aristatum grasses, middle of the 20th century, most Mexican species of Trisetum had been described. More recently. Morden and Valdés-Reyna (1983) described a new species, Т! curvisetum Morden & Valdés-Reyna, en- demic to Nuevo León, Mexico. Hernández-Torres and Koch (1988) have pub- lished the most recent revisionary study for all Mexican species of Trisetum, and they recognized 11 species: T. altijugum (E. Fourn.) Scribn.. T. cur- T. interrup- Lum, Torres, T pringlei, T. spicatum, T. viride, and T. virletu. Es- 0 visetum, T. deyeuxioides, T. filifolium, T. irazuense, T. kochianum Hern. pejo-Serna et al. (2000). as a part of the. floristic list of southern Mexico the presence of T. angustum Swa- Mexican monocotyledons, mentioned for Hen and J. pinetorum Swallen, both originally de- scribed from Guatemala. Other recent treatments of the genus are those of McVaugh (1983) for Nueva Galicia, Herrera-Arrieta (2001) for Durango, and Herrera and Rzedowski (2001) for Valle de Mexico. In Central America, Pohl (1980) in Flora Cos- taricensis mentioned four species: Trisetum deyeu- xioides. found also from Mexico to Central and South America, 7! irazuense, T. pringlei from Mex- ico to Central America, and J. tonduzii Hitche. known only from Costa Rica and Panama. Mc Vaugh (1983) and Hernández-Torres and Koch (1988) cit- ed T. viride from Guatemala. and Herndndez- Torres and Koch (1988) cited J. kochianum from Guate- mala and Costa Rica. The only subgeneric treatment of Trisetum in the Americas was done by Louis-Marie (1928—1920), who prepared a key to the subgenera, sections, and subsections. His Trisetum subg. Heterolytrum Lou- is-Marie sect. Anaulacoa Louis-Marie included: T. filifolium, T. interruptum (as T. disjunctum Louis- T. rosei Scribn. & Merr., T. Marie), T. irazuense, spicatum, T. viride (as T. paniculatum E. Fourn.), and T. virletii. In subsection Sphenophoidea Louis- Marie he placed Sphenopholis interrupta (as T. in- terruptum); in subsection Deschampsioidea Louis- Marie he placed J. palmeri; in subsection Graphephorum (Desv.) Louis-Marie he placed Pey- rischia. koelerioides [as T. altyugum (E. Fourn.) Scribn.. and J. pringlei; in subgenus /solytrum ouis-Marie he placed Peyritschia deyeuxioides Kunth) Finot (as Here. we describe and illustrate four new Mex- — Т. deyeuxioides) and T. viride. ican species of Trisetum (T. durangense Finot & P. M. Peterson, T. martha-gonzaleziae Р. M. Peterson & Finot, T. spellenbergii Soreng, Finot & P. M. Pe- terson, and T. ligulatum Finot & Zuloaga). and we discuss the systematics for Trisetum, Peyritschia, and Sphenopholis in Mexico and Central America. We also include a subgeneric classification of the species in Trisetum for Mexico and Central Amer- ica. MATERIAL AND METHODS This study is based on the examination of her- barium specimens from BAA, BAF, C. CHDIR, CONC, CR. P. QCA, US, S, SI. including the type specimens of most of the species studied. For an- atomical observations, hand cross sections. were made from the central portion of the blade below the flag leaf; and surface features were observed on the adaxial portion of the ligule. Abaxial epidermis preparations were made according to the method in Metcalfe (1960). Slides were observed on a Zeiss microscope at SL. KEY FOR DISTINGUISHING TRISETUM, SPHENOPHOLIS, AND PEYRITSCHIA la. Stamens 2: ше s linear, oe | -nerved: lemmas awnless or awned with a 2-lobate apex: palea tightly enc idee by ilie margins of the lem- ma (not gaping) Peyritschia Ib. Stamens 3: glumes or oblance Р li e first glume I- to -nerved; lemmas or entire or ^s lanc solate. OV ate-lanccolate. 3-nery mer second glume 3- to with 2 to 4 short awns at the ap 2-toothed: ds a not tightly enc lused by the mar- gins of the lemma (gaping). Perennials: disartic anaes between the flo- rets and above the glumes: upper glumes lanceolate to ov sue ырга n Trisetum 3a. Lemma apex opaque, the intermediate 1 s; awn inserted on the upper third of the lemma y я isetum subg. Тешип ш ТЕ le lax, + open, ovate to py- ramidal Trisetum sect. Trisetum 4b. Panicle narrow, spic iform — Trisetum sect. ма Annals of the 4 Missouri Botanical Garden 3b. Lemma apex hyaline, without nerves or Avena joi ie Kunth, Nov. Gen. Sp. 1: 148 1616, Tri- with both intermediate and marginal um elongatum (Kunth) Poir., Басус), 366. nerves extended beyond the apex as 817. Trisetum ee (Kunth) Kunth, Rev. four short awns; awn inserted on the an 1: 101. 1829. TYPE: Mexico. In scopulosis middle of the lemma prope urbem "hus 1833, aoe s.n. (holotype, risetum subg. Desc жое P!; isotype, US-3102206 2b. Annuals or perennials; disarticulation of flo- Calamagrostis н Buc tes. Pre lim. Rep. Surv. rets below the glumes: upper glumes oblar Texas 2. TYPE: U.S.A. Texas. [No type ma- ceolate to obovate Sphenspialis terial has bee sn loc rated, ak the type locality is presumed to be Texas, Buckley did not cite a loc ality TAXONOMIC. TREATMENTS n is did i mention a collector in his original pro- olog Sphenopholis Scribn., Rhodora 8: 142. 1906. Т kalli Se ‚ Bull. Torrey Bot. Club 11: 6, 1884. TYPE: Sphenopholis obtusata (Michx.) Scribn. Annuals or perennials, without rhizomes: culms 20-120 cm tall, glabrous. sparsely pubescent; ligule membranous; blades 1— «al sheaths glabrous or 12 mm wide, glabrous or sparsely pubescent, flat, soft. Inflorescence in panicles 5—25 cm long, lax 3-flowered; pedicels the glumes; glumes heteromorphic, usually shorter than and narrow. Spikelets 2- short and glabrous: disarticulation below the spikelet; first glume narrow, shorter than or as long as the second glume, linear-lanceolate, l- nerved; second glume obovate to oblanceolate, 3- to 5-nerved: lemmas smooth or scabrous, muticous or awned, 3- to 5-nerved, blunt to biaristate at the apex; awn, when present, borne on the upper third of the lemma: callus glabrous to subglabrous; palea 2-keeled, shorter than its lemma; lodicules 2, mem- branous, minutely dentate at the apex, 0.5 mm long; stamens 3, anthers 0.5-1 mm long: ovary gla- brous. Caryopsis with liquid endosperm. Sphenopholis is a small genus with five species from Canada to Mexico (Clayton & Renvoize. 1986). Erdman (1965) revised Sphenopholis and recognized four species in addition to the hybrid Sphenopholis Xpallens (Bichler) Seribn. [S. obtu- Scribn. X S. Two species are represented i pensylvanica | (L.) Mexico sata (Michx.) Hitche. |. and Central America. KEY TO THE SPECIES OF SPHENOPHOLIS IN MEXICO. AND CENTRAL AMERICA la. Lemmas awned, the awns 4—8 mm long: spike- lets 4-6 mm long 5. interrupta Ib. Lemmas unawned: spikelets 1. 5-3.6 mm n 2. S. obtusata la. Sphenopholis interrupta (Buckley) Scribn. subsp. interrupta, Rhodora 8: 145. 1906. Ba- sionym: Trisetum interruptum Buckley, Proc. Acad. Nat. Sci. Philadelphia 14: 100. 1862. TYPE: U.S.A. Middle Texas, Buckley s.n. (lec- totype, designated by Hitchcock (1935: 973), PH not seen.; isotype, US-fragment & photo ex PH!). ribn Senha hallii (Scribn.) Seribn.. pee 8(92): 1 46. ›. Fisetum inte و میا‎ | NR ED hallii (5с ib n. a Нисһе. TYPE: U.S 5 prairies Houston, 1872, Hall 799 Ыы US-72664!). Annual; culms 10—40 em tall, glabrous, mostly 3 nodes, the nodes glabrous. Leaf sheaths glabrous or sparsely pubescent, with one side extended up- ward, shorter than internodes; ligules 1 mm long, truncate, ciliate, blades 3-10 em X 1—4 mm, flat. Panicles 5-12 em X 4-15 mm, narrow. Spikelets 4-6 mm long, 2- or pilose on the dorsal surface: 3-flowered: rachilla sparsely pubescent; glumes 3.5-5 mm long, about equal in length and equaling or shorter than the spikelet, glabrous, scaberulous along the keel; first. glume linear-lanceolate, 3- nerved; second glume obovate to oblanceolate, 5- nerved; lemmas 3.5-5 mm long. lanceolate. gla- brous, 3- to 5-nerved, 2-aristulate at the apex, awned on the upper third of the lemma: awn 4-8 mm long, slightly twisted and twice geniculate; pa- ea 2-2.5 mm long. about half the length of the lemma, 2-nerved, the nerves scabrous, 2-dentate at the apex: callus glabrous to subglabrous; lodicules 0.5 mm long, slightly bilobate at the apex, the lobes Anthers about 0.5 mm rounded, obtuse to acute. long. Distribution. U.S.A. and Mexico. According to Gould and Moran (1981), Sphenopholis interrupta is the southwestern United widely distributed in States. MEXICO. Baja California: near 1 Apr. 1886, Specimens studied. U.S. Boundary, “Northern Lower California." C. R. Orcutt 1431 (US). KEY TO THE SUBSPECIES OF SPHENOPHOLIS INTERRUPTA S. interrupta subsp. interrupta la. Glumes glabrous ^ 5. interrupta subsp. californica Ib. Glumes pubescent Ib. Sphenopholis interrupta subsp. californi- ca (Vasey) Scribn., Rhodora 8: 146. 1906. Ba- sionym: Trisetum californicum Vasey, U.S. D. A. Div. Bot. Bull. 13(1): t. 46. 1892. Trisetum in- terruptum Buckley var. californicum (Vasey) Louis-Marie, Rhodora 30: 240. 1929, TYPE: Volume 91, Number 1 2004 Finot et al. 11 5 Peyritschia & Sphenopholis Mexico. Northern Lower California, C. К. Or- cutt 1437 (lectotype. designated by Hitchcock (1939: 552) and Hernández-Torres & Koch (1988: 80), US-868402!; MO- 2526390 not seen). isotype. Glumes pubescent, the trichomes 0.1—0.4 mm long. Distribution. lifornica was described for Northern Lower Califor- Sphenopholis interrupta subsp. ca- nia, and its distribution seems to be restricted to the type locality. Notes. Hitchcock (1939) designated the lecto- type for Trisetum californicum without explicitly cit- ing the herbarium. Hernández-Torres and Koch (1988) then specifically designated the US sheet as the lectotype. Comments. Though the systematic position of this taxon in Sphenopholis was proposed by Serib- ner (1906), subsequent treatments by Louis-Marie (1928-1929). Hitchcock. and Chase (1951). Erdman (1965) excluded this species from Sphe- and nopholis. According to Erdman (1965) further study is needed to clarify the generic placement of this species. On the basis of micromorphological char- acters of lemma epidermes of several genera in Aveninae, including Trisetum and typical species of Sphenopholis, Finot and Matthei (unpublished re- sults) support the generic position of this species in Sphenopholis. Louis-Marie (1928-1929) created Trisetum subsect. Sphenophoidea for species that disarticulate below the glumes and included in this T. interruptum, T. interruptum var. californicum, T. hallii, T. pensylvanicum (L) P. Beauv., T. palustre (Michx.) Torr.. lieve that all of these species belong in Sphenop- holis. and T. ludovicianum Vasey. We be- N . Sphenopholis obtusata (Michx.) Seribn., Rho- dora 8(92): 144. 1906. Basionym: Aira obtu- sata Michx.. Fl. Bor. Amer. 1: 62. 1803. Ai- ropsis pid (Michx.) Desv.. J. Bot. appl. 1 200 . Poa шам (Michx.) Link, Hoy: Berol. 1: E^ Agrostis 1 (Michx.) Steud., Nomencl. Bot. ed. 2. 1: 41. 1840. Koe- leria obtusata (Michx.) Trin. ex i» d., No- mencl. Bot. ed. 2. 1: 41. 1840. Reboulea ob- tusata (Michx.) A. Gray, Manual 591. 1848. Eatonia 1 (Michx.) A. Gray, Manual ed. 2, 558. 6. TYPE: U.S.A. Florida: Michaux s.n. duas pe. designated by Hitchcock (1908: 152), Pl. photo NY not seen & US! US-72667-fragment!). ; Isotype. Deser. Gram. 83. 1817. Koeleria Aira truncata Muhl., ) Torr. Fl. N. Middle United States . TYPE: U.S.A. Pennsylvania: е 127 (holotype, PH not seen; isotype, U Ф 1535792-fragment!). Koeleria 1 Nutt., Gen. N. Amer. Pl. 2: (Add. 2). 1818. : U.S.A. East Florida. J. Say s.n. (ho- da PH not seen, photo US!; isotype, US-frag- truncata er 1: 116. itt). канна gracilis з Revis. Gram. 2: 341, t. 84. 1630, illeg. TYPE: Michaux s.n. (holotype, B not seen; isotypes, 34 US- 76319- dang ex В!). Trisetum pts Trin., Mem. Acad. Imp. Sci. St.-Peters- urg, Ser. 6, Sci PME ]: 66. 1 аорта obtusata subs lob a (Trin.) Scribn.. Rhodora 8: 144. 1906. ee obtusata var. lobata (Trin.) Scribn. ex B. L. Rob., Rhodora 10: 65. 1908. TYPE: Ad flumen dc a (Red River), inna borealis holotype, LE-TRIN- 2437.01!). Eatonia density ra E. Fourn.. 1886. y 11. 1886. nom. inval. TYPE: U.S.A. Texas: Bear Feb. 1839, J. L. Berlandier 1617 (holotype, W not seen: e S, B not seen, LE-TRIN-1951.02!, US-72668- fra nt!). Eatonia pre var. robusta Vasey ex L. H. Dewey, Contr. U.S. Natl. Herb. 2: 544. 1894. аси гоню (Va- sey ex L. H. Dewey) Rydb., Bull. v Bot. Club 32: 602. 1905. Sphenopholis robusta em ex L. Н. . 6: 12. 1910. TYPE: ille, i Wallis s.n. (holotype, otype, NY not see Eatonia pene var. robusta 5 ex Rydb., Contr. U.S. Nall. Herb. 3(3): 190 5 nom. illeg. hom. TYPE: U.S.A. Nebraska: Mullen, P. A. Ry a erg 1807 (ho- зе, not located; 1 NY, Eatonia obtusata Mo 5 uem ex Rydb. & Shear. Bull. Div. Agrostol. U.S.D.A. 30. 1897. е уаг. Mes fo. purpurascens \ tydb. & Shear) Waterf., Rhodora 50: 93. 19. a TY i TSA Oklahoma: Palmer 404 (lec m Bus ock (1935: 956), US-868756!). Eatonia еи Seribn. & Merr.. n il. Div. Agrostol. U.S.D.A. 26: 6. 1900. Sphenopholis d subsp. Mr ear ы ribn. & Merr.) Scribn., Rhodora 8: 144 1906. Sphenopholis obtusata var. une ens (Scribn. & Merr.) Seribn. ex B. IL. Rob., Rhodora 10: 65. 1908. Sphenopholis 5 (Seribn. & Merr.) A Heller, Muhlenbergia 6: 12. 1910. Re Boule eres False Washita, designated by var. pubescens (Scribn. & T rr.) Farw.. id ». Michi- gan Acad. Sci. 17: 181. 1916. TYPE: n sissippi: Starkeville, 30 Apr. 1891, 5. i "Tay s.n. (holotype, US-72070!). W. Amer. Sei. 15: 50. 1906. Sphe- nopholis е (Suksd.) A. Heller, а Ө: 910. TYPE: U.S.A. Oregon: n Jalles on iiw River, 8 June 1897, r^ pese 1553 (isotypes. F not seen, GH not seen, ISC not NY not seen, UC not seen, US-72 т annua Suksd., seen, Annual: culms 20-120 em tall. Leaf sheaths gla- brous, scabrous. or finely pubescent: blades 8-15 cm X 3-12 mm, flat. Panicles 5-25 cm X 5-20 mm, narrow. Spikelets 1.5-3.6 mm long, 2- or 3- flowered: rachilla mostly glabrous; glumes 1—3(- 3.6) mm long. about equal in length, dimorphic, Annals of the Missouri Botanical Garden shorter than the spikelet, scaberulous especially along the keel: first. glume linear-lanceolate, 1- nerved; second glume conspicuously obovate, 3- nerved; lemmas 1.8-2.8 mm long, lanceolate. gla- brous to scaberulous, entire at the apex. unawned: callus glabrous; anthers 0.5—0.7 mm long. Anatomy and micromorphology. ligule apices with papillose cells and short, stiff trichomes; ligule epidermis composed of long cells with straight walls; stomata absent; macrohairs absent; prickles present. Foliar epidermis with long cells rectan- gular to fusiform, with sinuate lateral walls: short cells present in costal zones only, rectangular, with sinuous walls: trichomes absent; stomata present. MEXICO. ¢ Diaz, Specimens studied. Coahuila: Grande V Valley, е Hitchcock 5893. Di urango: city ar D u- rango & vicinity, June li 890. : digi 255 (US). 0 (US) Nuevo. León: Tiere rev, re vis ALS. ‘cd 5571 ‚ Oaxaca: Oaxaca . 12, 1 ug. 1910, LU pos (US). Puebla: Fane E E .20 ud . Nicolas s.n. (US); С. ae s.n. (US). 577 (US 1890, V. Palmer denda d Mexic. Pl. 2: koelerioides 109. 1880. (Peyr. E. Peyritschia E. Fourn., TYPE: Fourn. Peyritschia Perennials, with or without rhizomes; culms 20— 200 cm tall, mostly glabrous. Leaf sheaths glabrous or pubescent: blades flat: ligule membranous. Inflo- rescence in panicles narrow, contracted, spiciform or lax and somewhat open. Spikelets 2-flowered: rachilla glabrous to copiously pubescent: disartic- ulation above the glumes and between the florets: glumes isomorphic, 1-nerved, equal or subequal. as long as the spikelet or longer: lemmas bilobed at the apex, awned near the base or the middle of the lemma, rarely muticous or shortly mucronate near the tip (P. koelerioides mucronate or muticous): pa- lea tightly enclosed by the margins of the lemma (not gaping), 2-keeled; stamens 2; lodicules 2. membranous; ovary glabrous. Caryopses with liquid endosperm. KEY TO THE е OF PEYRITSCHIA IN MEXICO AND CENTRAL AMER | а. Lemma without а 17 awn, apex muticous o1 with a subapical тист 2. P. koelerioides Lemma with a dorsal awn, borne near the middle or upper third of the lemma, the awn geniculate and extended beyond the glumes. — 2a. Rachilla copiously pubescent, ER trio homes lor 2-3 mm long deyeuxioides 2a. s pubescent, the trichomes only 0.2— 0.8 mm long. 3a. C ulm is 5-12 cm tall: leaf blades 2.5-3.5 em long: panicles about 2.5 em long, 0.5 em wide, contracted, narrow P. humilis le af lade 8 5- w C ulms 20—200 em tall; poni narrow but 1 ‘ly open P. pringlei l. Peyritschia deyeuxioides (Kunth) Finot. Contr. U.S. Natl. Herb. 48: 478. 2003. Ba- sionym: Avena deyeuxioides Kunth, Nov. Gen. Sp. 1: 147. 1816. e E (Kunth) Poir., Encycl. Suppl. 5: 366. 1817. Trisetum deyeuxioides (Kunt и Kinik. Rev. Gram. 1: 102. 1829. i a triflora Nees, Linnaea 19: 691. 1847. : Mexico. "Ad | отой & Воп- pland s.n. (holotype, PI; isotypes, BM not seen, LE-PRIN-1913.05!, US-865589 fragment. ex PD. ripam. Lacus Tezcucensis,’ Avena tric hopodia J. Presl, Reliq. Haenk. 1: 254. 1830. TYPE: Mexico. Haenke s.n. (hok otype, PR not seen; T E-TRIN-1945.01a!, US-0865583 frag- ment ex Ad sa evoluta | К. Fo Bull. . Bot. France 24: 81. 18 Trisetum popu. (. i Hitchc., С U. s. “Кай, Herb. 17: 325. 1913 РЕ: Mex- ‚ Veracruz: Mirador, 1841, КОМ. тк 730 (le 'clobtype, абаке d by Hitchcock (1939: 557) and Hernández-Torres & Koch (1988: 78), US-207459!: isolypes, C not. seen, LE! MO-3056872!. P not seen). Trisetum var. Beal, d pr Scribn. ex Gras Ў e 374. . TYPE: Mexico. dn acán: dry hills near diei 'uaro, 19 Nov. 1891, G. Pringle 3950 а MSC not seen; isot H not seen, 3 not seen, МЕХ 3050871 not seen, P!, US-868407!). Bs ^yeuxioides 5 М. А! "у pes, MO- 2: nol seen, Perennial, without rhizomes: culms 50-180 cm tall, glabrous. Leaf sheaths glabrous or pubescent; ligule 1—3(—5) mm long, truncate, erose, glabrous: blades 0.54 mm wide. flat, narrow toward apex, glabrous or sparsely pubescent, sometimes densely pubescent. Panicles 10-35 1-5 ст, lax, some- what open and nodding. Spikelets 4.5-8 mm long, 2- or 3-flowered: rachilla copiously pubescent, the trichomes 2-3 mm long; glumes (3.5-)4-5.5(-8) mm long, equal, shorter or longer than the florets, linear; first glume with a green midnerve, purplish otherwise, the margins usually hyaline, the keel scabrous; lemmas 4-6 mm long. glabrous, terete, rounded in cross section, 5-nerved, nerves incon- spicuous, dorsally awned near the middle or upper third of the lemma: apex entire, bilobed or biden- tale: 4.5-8(-12) mm long. geniculate and twisted, purple, scabrous; callus short. pubescent, awn Volume 91, Number 1 2004 Finot et al. 7 1 8 a Peyritschia & Sphenopholis the trichomes 0.3-1 mm long: paleas 2.5-3.5 mm long; lodicules 0.5 mm long, linear, narrow, irreg- ularly lobed at the apex: anthers 1.3-1.7 mm long. Caryopses 1.8-2.5 mm long. Chromosome number. 2n = 14, 28 (Tateoka, 1962: Hernández-Torres & Koch, 1988). Anatomy. Leaf blades are slightly keeled in transverse section, abaxial ribs not well developed. furrows less than one half of the leaf thickness: vascular bundles of first and second order with ad- axial and abaxial girders, bundles of third order with adaxial and abaxial strands; marginal scleren- chyma present; bulliform cells large, conspicuous, in fan-shaped groups at the base of adaxial furrows: abaxial epidermis with long rectangular to fusiform intercostal cells; costal zone with cork cells and silica bodies in short rows or more rarely in cork cell-silica body pairs; prickles present. Distribution. Costa Rica, El Mexico (Aguascalientes, Chiapas, Chihuahua, Dis- Mexico to Ecuador Colombia. Salvador, Guatemala, Honduras. trito Federal. Durango, Guerrero, Guanajuato, Hi- dalgo, Jalisco, México, Michoacan, Morelos, Oaxa- ca, Puebla, Veracruz. Zacatecas). Nicaragua, Panama, Venezuela | (Pohl & Davidse, 1994: Finot, 2003a) in humid prairies and open woods, 1000—2000 m. Notes. Hitchcock (1939) designated the lecto- type for Deyeuxia evoluta without explicitly citing the herbarium. Hernández-Torres and Koch (1988) then specifically designated the US sheet as the lectotype Queretaro, San Luis Potosí, Comments. Based on 25 morphological char- acters, Koch (1979) pointed out that P. deyeuxioides shows an intermediate position between Peyritschia and Trisetum. However, the androecium is com- posed of only two stamens, the glumes are l- nerved, the lemma is terete and dorsally awned near the middle, the apex of the lemma is bilobed, and the palea is tightly enclosed by the margins of the lemma. These characters indicate that this spe- cies should be placed in Peyritschia. Specimens studied. COSTA RICA. Cartago: Cordil- lera Central, Volcán Irazú. 2 km NW к, reg: 19 Nov. 1968. Pohl & Davidse 11449 (CR). MEXICO. Chi- huahua: 19.6 km W of Balleza & 74.2 km E и Guacho- chi, 18 Sep. 1991, Peterson et al. 10752 (US); 24.2 km 5 of San Juanito & 4.8 km N of € d on MEX 127, 10 Sep. 1989. P eterson et al. 8007 (US); 122.5 km W of La Junta d y зк hic in (ame Nacional Cascada . 1989, Peterson & King 8231 (US); of = 200. 13 Oct. 1887, Pringle 1432 (US); Sierra Madre Oc c ide p Parque Natural 1 a del Cobre, 13.6 mi. 1 Bufa on road toward 5 31 Oct. 1995, 9 ор el al. 13575 (US 3. (os: иша: Sierra Madre Oriental, 5 mi. W of Chapultepec on — canyons 2 Nee cutoff road between Hwy. 54 & 57, 23 mi. S of Saltillo, 16 Oct. 1995, Peterson & Knowles 13276 (US); ca. 5 km Е of Salltillo (Las Palapas) up camino de Quatro, 20 Sep. 2003. Peterson et al. 17868 (CIIDIR, US). Distrito Fed- eral: Desierto de Los Leones, 19 Oct. pi Tateoka 1152 (US). кош 2 km of San Miguel de La Michilia. 24 Sep. 1991. Powis et al. 10913 (US); 16.1 km NW of MEX 40 on logging road to Santo Domingo, 5 Oct. 1989, Peterson & King 8288 (US); 20 mi. SE of ео on road to Charcos, 7 Nov. 1995, Peterson et al. 136. Hida mat bogey us of Lago 2 a below ш 24 July 1947, Moore Jr. 3466 ). México: Valle de Mexico, 10 үте 1901, Pringle 051 (P). Shaffner 15 (P); prope Tacubaya, Shaffner 319 (P). Michoacán: 15.5 km SE of Zacapu on MEX 15 km toward Quiroga. | Oct. 1991. Peterson et al. 10994 (US). Morelia: San Miguel, 8 Aug. 1912, Arsénes 8645 (P). Oaxaca: on weep Road 19 km from Ti . 1963, McKee 10869 (P). Querétaro: 16 mi. E of Landa, 12 Dec. 1900, Smee & Johnston s.n. (US). San Luis Potosi: between Puerta Huerta and Rioverde in the Sierra de Alvarez, IT Sep. 1954. Sohns 1177 (US); near Puerta Huerta in the Sierra de Alvarez. 4 Sep. 1954, Sohns 1034 (US). 2. Peyritschia koelerioides (Peyr.) E. Mexic. Pl. 2: 1886. Aira koelerioides Linnaea 30: 5. 1859. Deschampsia koe- lerioides (Peyr.) Benth., J. Linn. Soc., Bot. 19: 96. 1881. TYPE: Mexico. México: Volcan de Toluca. 8800 ft., 1846, C. Heller 311 (holo- type. P not seen; isotype, US-1647945-frag- ment ex P-DRAKE)). Fourn., Peyr., Е . E. Fourn., Mexic. Pl. 2: 111. 1886. tum. altijugum Е. Tourn. | pg dora 8/89): 89. 1906. TYPE: Mex SJ 8 xico. Veracruz: in monte Orizabensi, 5 ft. T. M. Lieb- not seen; isotype, US- mann 603 (holotype, P 5!). culms 25-85 cm tall, glabrous; nodes 2 or 3, glabrous. Leaf sheaths glabrous, rarely pubescent, shorter than the inter- nodes; ligule 1-3 mm long, truncate, erose; blades 1.5-4 mm wide, usually glabrous, flat. Panicles 7— 20 cm X 4-8(-10) mm, spiciform: rachis and ped- icels scabrous. Spikelets 3-5 mm long. 2-flowered: Perennial, rhizomes absent: rachilla glabrous to sparsely pubescent, the tri- chomes less than 0.5 mm long; glumes 3—5 mm long, lanceolate to ovate-lanceolate, equal, longer than the florets, 1-nerved; lemmas 2.5—4 mm long. glabrous, terete, dorsally rounded, 5-nerved; apex унет the lobes short, obtuse, muticous or with a mucro about 1 mm long or less borne near the apex: callus glabrous or short pubescent, trichomes less than 0.5 mm long; paleas 2.4-3.8 mm long: anthers 0.5-1.3 mm long. Caryopses 1.5-3.5 mm long. Chromosome number. 2n = 28 (Tateoka, 1962: Hernández-Torres & Koch, 1988). Annals of the Missouri Botanical Garden Micromorphology. — Ligule apices have papillate cells and short trichomes, the epidermis with more or less rectangular long cells, hooks, and trichomes: no stomata were seen, Distribution, Southern Mexico (México, Mi- choacán, Oaxaca, Veracruz) and Guatemala (Pohl & Davidse, 1994; Finot, 2003a). 2400-4300 m. ‚решили studied. MEXICO. México: 20 km al NE Texcoco, sobre la carretera a ME 10 Oct 1965. Rzedowski 21417 (US); Sic | Oct. 1892, Pringle 4306 (U М); Amecamec a, carretera al Paso de Cortés, 20 a la carretera a a 18 Oct. 1970, S of Patzcuaro, 20-25 King & Soderstrom 5159 (US); 20 km NE de pt 0. sobre la carretera a С alpulalpán. no date, Rze- dowski s.n. ). Oaxaca: Sierra de San Felipe. Aug. 1894, Pringle 477 75 (P). Puebla: San Nicolás de los Ran- chos, declive E del Paso de Cortes, 3.5 km NE entronque de carretera a Amecameca y camino entre Popoc atepetl e ra Las Cruces, м 90 Eus ESE de Amecameca, 3 1976, Koch 73309 3. Peyritschia humilis (Louis-Marie) Finot. Contr. U.S. Natl. Herb. 478. 2003. Ba- sionym: Trisetum humile Louis-Marie, Rhodora 30: 244. 1929. Deveuxia gracilis E. Fourn.. Mexic. Pl. 2: 106. 1886, nom. illeg.. non De- yeuxta gracilis Wedd. 1875. TYPE: Mexico. Mirador, К M. Liebmann 602 (holotype. Cl: isotypes, US-3048345 Haun.!). fragment ex hb. Perennial: culms 5—12 cm tall. Leaf sheaths flat- tened; ligule membranous. short, laciniate: blades 2.5-3.5 cm long. Panicles 2.5 X 0.5 em. contract- ed, narrow, exserted. Spikelets 4 mm long, 2-flow- ered; pedicels glabrous; rachilla 0.5 mm long. pi- lose, the trichomes less than 0.8 mm long: glumes і mm long, equal, lanceolate, longer than the spikelet, slightly scabrous on the keel, acute at the apex: lemmas 3-3.5 mm long, terete, awned. the margins involute toward the base, bilobed at the apex, the lobes obtuse: awn 4—5 mm long, inserted from the middle to the lower third on the dorsal twisted and callus obtuse, surface, geniculate: sparsely pilose, trichomes ca. 0.2 mm long: paleas shorter than the lemma and enclosed by the mar- gins of the lemma. Caryopses ca. 2 mm long. Comments. The type of Deyeuxia gracilis E. Fourn. conserved in Copenhagen (C) includes a complete plant, 12 cm tall, and a fragment of the panicle, identified by Hitehcock and Chase as Tri- Richt. identification. setum spicatum (L.) K. The isotype in US bears the Hernández-Torres and Koch within J. spicatum., followed this criterion. The mor- same (1988). who synonymized the species phology of the spikelets confirms its placement within Peyritschia: spikelets 4 mm long, 2-flowered: glumes equal, l-nerved, longer than the spikelet; lemma with a bilobed apex, the awn geniculate and twisted, inserted near the middle of the lemma: cal- lus with short trichomes; rachilla sparsely pilose; stamens 2; ovary glabrous. Trisetum humile Louis-Marie was a new name for the illegitimate Deyeuxia gracilis E. Fourn. 1886. non Weddel 1875, and therefore we use it as the basionym for the new combination in Peyritschia. 4. Peyritschia pringlei (Scribn.) S. D. Koch, Tax- on 28: 233. 1979. Basionym: Deschampsia pringlei Scribn., Proc. Acad. Nat. Sci. Phila- delphia 43: 300. 1891. Trisetum kochianum Hern. Phytologia 61: 4. 1987 TYPE: Mexico. Chihuahua: wet places, pine plains, base of Sierra Madre, 7 Oct. 1987, C. G. Pringle 1429 (holotype, US-747292!; iso- types, LL not seen, MEXU not seen, MICH not seen, US-867629!, US-821538!. VT not seen). Torres, Perennial, without rhizomes; culms 20-200 em tall. Leaf sheaths glabrous, rarely pubescent: ligule 0.5—4 mm long, truncate or rounded at the apex; 5-20 X 1-34) em, narrow but relatively open. Spike- blades 5-15 em X 1.5-4 mm, flat. Panicles lets 4-5.5 mm long, 2- or 3-flowered: rachilla pu- bescent, the trichomes 0.2-0.8 mm long: glumes 4— 5.3 mm long, lanceolate, as long as the florets or longer. apex acute: lemmas 3-4.3 mm long, gla- brous, terete, bilobed at the apex. rounded in cross section, awned near the base; awn 4.5-7 mm long, geniculate and twisted, exserted: palea 2.3-3 mm long, shorter than its lemma; anthers 0.8-1 mm long. Caryopses 1.5-1.7 mm long. fusiform- cylin- drie. Inatomy and micromorphology. Ligule apices composed only of papillate cells, epidermal long cells rectangular to fusiform, with few hooks and no trichomes: no stomata were seen. Leaf blades in transverse section without a keel, without well-de- veloped ribs, furrows less than one half of the leaf thickness: all vascular bundles, except the marginal ones, with both adaxial and abaxial girders; mar- ginal sclerenchyma absent; abaxial epidermis with rectangular long cells with straight side walls. cos- tal cork cells and silica bodies present. prickle hairs present. Distribution. Mexico (Aguascalientes, Chihua- hua, Chiapas, Durango, Guanajuato, Hidalgo, Jalis- San Luis Potosí. So- Rica, Venezuela, Ecuador (Espejo-Serna et al., 2000: Fi- ». México, Puebla, Querétaro. nora, Tlaxacala). Guatemala. Costa Volume 91, Number 1 2004 Finot et al. sven Peyritschia & Sphenopholis not, 2003a). The documentation of the presence of this species in Venezuela and Ecuador provides new records for the distribution of Peyritschia prin- glei. It was collected by Peterson & Judziewicz 9302, in the Provincia de Bolivar, ca. 3000 m, in the Andes of Ecuador, and by Peterson & Judzie- wicz 9477 in Provincia de Loja, Ecuador. Lasser found the species in Mérida, Venezuela. in 1942 (Lasser 431). Koch (1988) this species also occurs in Guatemala. Elevations range from 1600 to 3270 m. According to Hernández-Torres and Specimens studied. COSTA RICA. Cartago: Cordil- lera Central, lower slopes of Volcán Irazú. 1 km below 4 Juan A р 8 Nov. 1968. а & Davidse 11416 ( ECUADOR. Bolivar: 14.5 5 of Guaranda on ы to Río es 24 May 1990, us & Judziewicz 9302 (US). Loja: 6 km W of Colaisaca on road to Sozoranga, 2 June 1990, Peterson & Judziewicz 9477 (US). GUATE- MALA. Quezaltenango: 19 June 1954, Koninck 137 (US). MEXICO. Chihuahua: 85.4 km W of Balleza and 8.8 km E of Guachochi. 19 Sep. 1991, Peterson et al. 10786 (US); 38.6 km SW of La Junta « ca. 70.8 km of Creel at t Pas so Arroyo Ancho crossing, 24 Sep. 1988, Peterson & Annable 5848 (US) Miñaca, 13 Oct. 1910, 5). Durango: 45 km W of Durango on Hwy. 40. toward 5 Salt to. 2 Oct. `~ PA шын ‚ 1995. Peterson et al. 13679 (US); 24 mi. NE of El Salto on Mani 40 ud Durango. a Oct. 1995, Peterson et al. 13471 (US); W of Ciudad Durango, У mi. W of E Salto, s Aug. 19: 58. Reeder & Reeder 3124 (US): 4 mi. E of El Salto, 1 Oct. 1953, ке « кы: 2561 (US): 20 mi. S of Mezquital. 11 Sep. 2 chez-Alvarado 17729 (CHDIR. 5 M e San Felipe. 27 Oct. ТА Sohn 0 (US). Hidal- go: Pachuca. 6-7 Sep. 1910, Hitchcock бт 30 (US): Hitch- cock 6737 (US). Jalisco: Sierra Cacoma. Y mi. NW of Los Volcanes on road toward Mascota. 15 Nov. 1995, Peterson 13724 (US); Tepatitlán-Guadalajara, 26 Sen, 1946. Her- nández X., Ruppert & Guevara 2694 (US); Zempoala, 15 km N of Ciudad Sahagán & 1 km NE of л -mpoala, sobre la carretera a Sta. Maria Tecajete. 26 Aug. 1977. Koch 77179 (US); Mineral del Chico, Parque Natural El Chico, 17 km al N de la carretera Pac 'huca-Zac xL 4 Dec. : Toluca. 13 5 1910, 1 Marcos. sien em- 977. Koch 77532 (US Нисћсос i 6889 (US). bankment, 29 Aug. El к. А 10 km al E de ye 19?09' N, . Tenorio 15165 (US). Querétaro: erro Zamorano, Min: с Pa 13 Nov. 1971. Rze- dowski & ` MeVaugh 473 (US). = Luis Pavia: 15-18 July 1910, Hitchcock 5685 (US): Cañón del Lobo, Sierra de San Miguelito, Sohns 1/62 үа Puerta Huerta i in the 1 Sep. 1954. Sohns 1019 (US cala: ca. ‚ NE of Tlaxco. 23 Sep. 1953. Sohns 597 (US). VENEZUEL A. pa m Banda. near Mérida. 22 Sep. 1942, Lasser 431 ( i mas Sierra de Alvarez, m Pers.. Pl. 1: 97. 1805. Trisetarium Poir, Encycl. Suppl. 5: 365. 1817. nom. su- perfl. Acrospelion Besser ex Schult. & Schult. f.. Syst. Veg. Mant. 3: 526, 1827. nom. illeg. superfl. Rebentischia Opiz, Lotos 4: 104. 1854, non P. A. Karsten 1869. TYPE: Ess. Syn. nom. superfl., Trisetum rd (L.) P. m à togr. 88, 153, t. 18. f. 1. Agros- Rupestrina Prov., Fl. Canad.: 689. 1862. TYPE: Rupestri- na pubescens Prov. [= Trisetum spicatum (L.) Richt.] Perennials, caespitose, sometimes shortly rhizo- matous; culms 5-300 cm tall, erect to geniculate at base. glabrous or pubescent. Leaf sheaths gla- brous or pubescent, longer or shorter than the in- ternodes; blades flat, conduplicate. convolute or in- soft, Inflorescence in panicles contracted or open, spi- volute, rarely rigid; ligule membranous. ciform, ovate, or pyramidal; the rachis glabrous. scabrous, or pubescent. Spikelets (1)2- to 6-flow- ered, short pedicellate: rachilla pubescent or gla- brous, usually prolonged beyond the upper floret: disarticulation above the glumes and between the florets; glumes heteromorphic. lanceolate to ovate- lanceolate, equal or unequal, first glume l- to 3- nerved, usually shorter and narrower than the sec- ond. second glume 3- to 5-nerved: lemmas lanceolate. (3)5(7)-nerved, usually awned or muti- cous, with apex and margins hyaline, glabrous or pubescent, slightly keeled and compressed, rarely terete: apex with 2 to 4 short awns, entire, or 2- toothed: central awn from the upper third. rarely the middle, of the subapical portion of the lemma: awn exserted, geniculate or merely divaricate: cal- lus short pilose; palea not tightly enclosed by the margins of the lemma (gaping). 2-keeled, hyaline. usually shorter than the lemma: stamens 3, anthers T 34.5 mm long: lodicules 2, membranous: ovary ИЕ or with short and shining trichomes near the apex: endosperm solid or liquid, soft or hard. Caryopses compressed, soft; hylum short, puncti- form. Basic chromosome number x = 7. Our treatment of the species that occur in Mexico and Central America Comments. subgeneric includes two subgenera: Trisetum subg. Trisetum and 7! subg. Deschampsioidea, and two sections in subgenus Trisetum: section Trisetum and section Trisetaera (Appendix 1). In the following key we have indicated what subgenus and/or section most species occur in (see leads 5a, Sb, ба, and 8a). Ккү TO SPECIES OF TRISETUM IN MEXICO AND CENTRAL AMERICA la. Lemmas with the apex entire, slightly bidentate, or irregularly dentate; nerves of the lemma not prolonged bevond the apex: lemma muticous or with a short subapical awn (but awned in T. filifolium var. aristatum). Annals of the Missouri Botanical Garden 2а. Glumes o ry е in length, shorter than the spikelet, covering nearly half of the spikelet length; spikelets I- to 3-flowered: lemma with the apex obtuse to irregul: wly dentate; leaf blades flat or some- times a or involute, never filiform; lodicules lance salate, acute at the apex. Ja. Panicles 4-12 em wide, open, 5 the branches capillary and flexuous: first glume 1. 42. 6 mm long: second glume 24.5 mm long T. pringlei ЗЬ. Panicles 0.6-1 em wide, narrow and densely flowered; the branches с lose ly appre sed; first glume (2.7—)3—4 mm long: second glume 4—5.8 mm long LE aiam 2b. Glumes 5 10 equal і in length, shoul two-thirds as long as the spikelet or as long as the spikelet spikelets 2- or 3-flowered: lemma with the apex slightly bidentate to entire; leaf blades involute. filiform: lodicules line ar, slightly bilobate at the apex. 4a. Lemma muticous or with a short subapical mucro 0.2-0.5 mm long За. T. filifolium var. шш 4b. Lemma awned on the upper third: the awn 3-5 mm long, geniculate 3b. T. Аш ж var. aristatum Lemmas bidentate at the apex; inte бей. imd — nerves or only intermediate nerves. prolon beyond the apex as 2 to 4 apical setae; apical teeth some mer wil nerves; lemma aristate, the awn we vell deve sloped. borne on the upper third, middle, or lower third of the lemma. not 1 5 da. Awn always inserted on the upper third of the lemma (sometimes ne arly in this Prim in J. palmeri); glumes unequal, the first glume shorter and narrower than the second la apex of the lemma with two setae formed by the diis ction of the intermediate nerves; lemma + keeled in cross section: palea е. gaping (Trisetum subg. Trisetum). 6a. Panicles 2-15 em ide lax. mostly open or narrow, со ovary pubescent ог glabrous: culms glabrous below the inflorese ‘ence (Trisetum sect. Trisetum). Ta. Lemmas (2.5-)2.74.3 mm long, smooth to slightly scabrous; apex of the lemma no ntate; dorsal awn delicate, curve "s first glume 1.7-2.5 mm long; sec i glume 2.8-3. po n long: lodicules truncate at the apex; endemic to Nuevo León, Mexic T curvisetum Tb. Lemmas 4.3-6 mm long, strongly scabrous; apex of | в жене biaristulate; Eo awn not delicate, geniculate: first slime 2.5-5 mm ones sec e glume 3.5—6.5 mm long: lodicules lobulate at the apex: in. central Mexico to Pe . T. irazuense Ob. Panicles 0.5—1.3(-2) em wide, spiciform or subspic iform; ovary glabrous; culms a ee nt below the inflorescence. За. Lemmas glabrous; glumes se die on the ke E not ciliate; awn slightly scabrous; paleas gem on the kee ili (Trisetum sect. Trisetaera Glumes shorter than the spike let, 1 05 de first l-nerved, the second 3-nerve ес. ligule 0.5-1 mm long; plant not rhizomatoi T. spicatum 9b. G lumes as long as the spi нет Mie ан (1)3-nerved, the second (3)5- -ne I ligule —4 mm long: plants rhizomatot T. ligulatum b. Lemmas ss scent; glumes se 1 or ciliate on the keel; awn sc 'abrous or lie: рага is ш r ciliate on the keels. 10 Cs c E on the keel and slightly shorter to as long as or longer than the spikelet; first glume 3.7-6 mm long, ovate-lanceolate; second кн 1.5—6.5 mm long. ovate- lane со to ovate; le mma awns plumose; paleas ciliate on the keels 9. T. rosei 10b. Glumes somewhat scabrous on the keel and shorter than the spikelet (about 24 the length): first glume 3-3.2 mm long, linear-lanceolate: second glume about 4 mm long, line ur- абсеоіаќа to lanceolate; lemma awns not plumose, but ciliate near the base paleas scabrous on the keels 6. T. 5 5b. Awn inserted on the lower third or near the middle of the lemma: glumes subequal, nearly as long as 1 ıe spikelet or longer; lemma with apical teeth without prolongation of the intermediate nerves or with 2 to | setae; lemma rounded in cross section; palea sometimes enclosed by the margins of the lemma, not pias (Trisetum subg. Deschampsioidea ). lla. Spikelets 2-flowere ы panicles narrow, subspiciform to spiciform, 0.6-2(4) em wide. 12a. Glumes equal in length and longer than the spikelet: spikelets 7.5-9 mm long: re mmas bidentate at the apex; panicles 1.5-2(4) em wide, a with purple and gold; ovary with trichomes at the apex 11. J. macibargensaleñas 12b. Glumes subequal, shorter than the spikelet; spikelets 6.5-7 mm long; lemmas with four setae at the apex; panicles 0.6-1 em wide, green: ovary glabrous at the a pex 13. T. pinetorum IBI ee 2- to 6-flowered; panicles open. lax, (1—)2—20 em wide. За. Culms 150-300 em tall, bambusiform, 5-7(-10) mm diam. below: lemma apex 2- or 4- — or -toothed, sometimes appearing almost entire, the lateral nerves do not contain s (vascular tissue 17. T. virletii 13b. C m — 50 em tall, herbaceous, usu: Шу less than 4 mm diam. Бе low: lemma apex 2- lobed, or 4-awned due to the projection of intermediate and marginal nerves (setae) beyond the apex. Ida. Lemmas dorsally hirsute below the awn insertion: ovary with trichomes at the apex T. durangense 14b. Lemmas glabrous or dorsally scabrous: ovary glabrous, rare ‘ly pilose (see T. martha- gonzaleziae). лм, Volume 91, Number 1 2004 Finot et al. 11 Trisetum, Peyritschia & Sphenopholis 15a. Lemma apex 2-lobed and 4-awned as teeth or setae l6a. Lemmatal awns borne above the middle of ilie lemma. Та. Lemma apex with four setae, the setae 0.6-1 mm 8 paleas slightly shorter than the lemma; callus trichomes 1.2-1.7 mm long — БЕРЕРИ 1 tonduzii 17b. Lemma apex sibus p setae; є baleen as long as or ni langer than the lemma; callus trichomes 0.5-1 mm long or les lob. nig awns borne near the aiddi or on lower М or 4 o 18 11 Ligules 3-6 lemma: L poar 18b. tric ae 8 1.5- 15b. Lemma apex 2-lobed 1 any additional awns or setae. 19а. Spikelets (4)5- to 6-flowered __ I9b. Spikelets 2- or 3-flower 20a. Ligules 3-6 mm long: rachilla trichomes 0.5—1(-1.4) mm long 2 3) mm he ава shorter than the lemma: rachilla 16. vd daban X the lemma. bre icu as long as or slightly longer than the 5 ; 2. T. palmeri | long ꝛñ ñ— T. viride 14. T. spellenbergii «d. T. palmert 20b. L icula s 0.5-1(— 2)n mm » long rac hills trichomes 1.5-3 mm long. 21а Glumes 7.5-9 m cles 1.5— 21b. iele 52-8 cm wide. open or somewhat mtract 1. Trisetum angustum Swallen, Phytologia 4(7): 423. 1953. TYPE: Marcos: between San Sebastián and summit of Volcán Guatemala. San Tajumulco, on top of dry ridge in pine forest. 13 Feb. 1940, J. A. Steyermark 35525 (holo- type, F-1040546!: US-2236479 frag- ment & photo ex F!). isotype, Perennial; culms 12-24 cm tall, glabrous, dense- ly caespitose. Leaf sheaths glabrous or pubescent below; ligule 1.5-2.2 mm long: blades 3-8 em | mm, flat. glabrous or sparsely pilose. Panicles 1-11 X 0.6-1 cm. branches closely appressed. Spikelets 4.6-0 mm narrow, densely flowered. the long, 2-flowered; rachilla sparsely pubescent, the trichomes about 0.6 mm long; glumes shorter than first glume (2.7-)3—4 mm long, l-nerved; the second glume 4—5.8 mm long, the spikelet, unequal; 3-nerved: lemmas (lowest) 4-4.8 mm long, sca- brous: apex irregularly toothed to 4-dentate, muti- cous or with a short subapical mucro; the mucro up to 0.5 mm long; callus obtuse, short pubescent, the trichomes 0.2-0.6 mm long; anthers 0.8-0.9 mm long. Caryopses ca. 2.1 mm long: endosperm solid, hard. Distribution. This species was described from a Guatemalan specimen by Swallen (1953). A spec- imen from Chiapas, Mexico (Matuda 2542a). was cited in Pohl and Davidse (1994): however, we were not able to study this. This species was not cited for Mexico by Hernández-Torres and Koch (1988 but was included in Espejo-Serna et al. (2000). — Comments. The above description ts based on G lumes 37 mm x long; 1 unequal, shorter than the и let: n long. equal, longer than the spikelet: pani- A) em wide, contracted, subspic iform: ovary pilose ite: 11. T. martha- has aleziae ee 16. T. viride the study of type material and includes the descrip- tion for Trisetum angustum given by Pohl and Da- vidse (1994), who also studied the holotype. This species appears closely related to T. pringlei. Pohl and Davidse (1994) suggested that T angustum might be better treated as a variety of J. pringlei since these species are morphologically very simi- lar. Both species have obtuse lemma apices and lack well-developed awns. These characteristics do not correspond well with the generic characteristics of Trisetum. Therefore, we consider the systematic position of these two species to be tentative or pro- visional. We have placed T. angustum and T. prin- glei in Trisetum subg. Trisetum but have not as- signed them to a section (see Appendix 1). 2. Trisetum curvisetum Morden & Valdés-Rey- na, Brittonia 35: 375. 1983. TYPE: Mexico. Nuevo León: Zaragoza, Sierra Madre Oriental. Ejido La Encantada, sitio La Tinaja, 1 km N del aserradero, ca. 23°55'N, 99°45’ W, 2830 m. 25 July 1981, Valdés-Reyna & Capó VR-1419 (holotype, TAES not seen; isotypes, ANSM not ¿NC MEXU not seen, MO- TEX not XALU not seen, ENCB not seen, 3102085 not seen, US-2978399!. NY not seen, seen, UT not seen, seen). Perennial, with short rhizomes; culms 50-150 em tall, glabrous, weak: nodes 4, glabrous. dark. Leaf sheaths shorter than the internodes, glabrous to sparsely pubescent or only ciliate on the mar- gins; ligule 1.1-2.5 mm long, membranous, oval. Annals o Missouri Botanical Garden ciliate or denticulate; blades (5-)9-20 em X 1.5- 8.5 mm, flat, glabrous or sparsely pubescent on the adaxial surface. Panicles 11—30(—36) х ax, mostly open, pyramidal, paucispiculate, rachis 2-15 cm. glabrous: pedicels 0.5—4.5 mm long, scabrous to- ward the apex; the branches flexuous, in verticels of five, ascending. widely spaced below becoming crowded near the apex. Spikelets 3-5.2 mm long, (1)2- or 3-flowered: rachilla 1-1.8 mm, glabrous to scabrous, sometimes with a few trichomes at the apex; glumes shorter than the spikelet, unequal, margins hyaline, apex acute, scabrous on the keel: first glume 1.7-2.5 mm long. linear-lanceolate, at- tenuate, I-nerved; second glume 2.8-3.8 mm long, lemmas (2.5—)2.7—4.3 mm long, lanceolate, laterally compressed, slightly ovate-lanceolate, 3-nerved; keeled, awned, 5-nerved, the nerves conspicuous, glabrous to scabrous, slightly bidentate at the apex or the apex entire, the margins and apex hyaline: awn 3—5.5 mm long. borne on the upper М or Y. less than 1 mm from the apex, flexuous, delicate, curved, scabrous; callus obtuse, glabrous or sparse- ly pubescent, the trichomes ca. 0.1 mm long: paleas 2-3.1 mm long, hyaline, shorter than the lemma. 2-nerved, the nerves scabrous, acute at the apex: 0.2-0.3 mm He an- lodicules oblong truncate, or 2 short, thers 0.5-1 mm long; ovary with 1 curved and shining trichomes on the apex. Cary- opses 1.7-2.2 X 0.4-0.6 mm; endosperm soft. Chromosome number. 2n res & Koch, 1988). Anatomy and micromorpholog y. — 28 (Hernández-Tor- Ligule apices composed of papillose cells: trichomes absent: lig- ule epidermis more or less rectangular, short, with straight walls; prickles and macrohairs absent. Leaf sections flat; keeled and well Жан adaxial ribs absent; bulliform cells in fan-shaped groups between vascular bundles: eat epidermis com- posed of fusiform long cells, with straight walls: prickles present: short cells present both in costal and intercostal zones; stomata present only in ab- axial epidermis. Distribution. Endemic Mexico, known only from Nuevo León, 2100-2500 m. Comments, Although the occurrence of oblong truncate lodicules in Trisetum curvisetum is unusual in the genus in comparison with the typical form (lobulate to irregularly lobed or toothed) in the ge- 1983), characters seem to pose questions as to the generic nus (Morden & Valdés-Reyna, no other position of this species. The presence of short, curved, and shining trichomes on the apex of the ovary and caryopsis confirms its generic. position. The habit of the plant, the structure of the panicle, the morphology of the spikelets, the presence of trichomes on the ovary, and caryopsis shape all suggest an affinity with 77 cernuum. арР studied. MEXICO. Nuevo León: Zarago- чта Madre Oriental, 7 7 km SE antes de La Encan- Zaragoza-Ejido La Ene antada, 25 July 1981. J. Valdés-Reyna & M. Сард VR- 1417 (US-297 a Paratype) ca. 15 mi. SW of ramberri, 24702'32.2"N, ‚ 20 Sep. ~ se et al. 16719 (/ I i: DUC US): ca. 17 mi. of des rri, 20 Sep. 2002, Peterson et al. 16723 ( peed ‘ONC, US): Sitio La Encantada- he Tinaja on road to La 1 1 23°53’ 14.4" N, 99°47'44.2"W, 21 Sep. 2002. Pe- terson et al. 16764 (ANSM, CONC. US) N = H a За. Trisetum filifolium Scribn. ex Beal var. fili- 2: 375. 1896. TYPE: Mexico. Chihuahua: cool slopes of the Sierra Madre. 9000 ft., 3 Oct. 1887. C. G. Pringle 1431 (holotype, US -8177 3 CM not seen, US-825600!). folium, Grasses N. Amer. isotypes, Perennial, caespitose, rhizomes absent; culms 30-60 em tall, scaberulous, o glabrous. Leaf sheaths oo г pubescent: ligule (0.2—)1—3.5 mm long. ШЕ idee a truncate, glabrous or with short trichomes; blades 5-22 cm X 0.5-2 mm. involute. filiform, curved and flexuous, glabrous or slightly scabrous. Panicles 12-15 X 2—3 сш, lax. somewhat open, narrow, exserted; the branches semiverticil- late, the half-whorls with 3 to 7(8) branches. the lower ones 4-6 cm long; rachis glabrous. Spikelets 3.5-6 mm long, (1)2- or 3-flowered: pedicels sca- brous: rachilla 1—1.2 mm long, with trichomes са. 1.5 mm long. the trichomes longer at the apex: glumes subequal, shorter, slightly longer, or about as long as the spikelet: first glume 2.5-6 mm long, 1(3)-nerved, second glume 3-7 mm long. 3-nerved: lower floret 3.5 mm long: lemmas 3.5—5 mm long, obscurely 5-nerved, strongly scabrous, sometimes scabrous-pubescent, terete, the margins involute enclosing the palea: apex entire or shortly bilobed, each lobe with two short teeth, muticous or with a short subapical muero: mucro 0.2-0.5 mm long: callus pubescent or subglabrous, obtuse, trichomes са. 0.2 mm long: paleas 34.5 mm long. shorter than the lemma, enclosed by the involute margins of the lemma (not gaping), 2-keeled, the keels well apart, ciliate or conspicuously scabrous: lodicules with acute apex and a lateral lobe below the mid- dle, 0.4—0.5 mm long: anthers 1-2 mm long: ovary glabrous. Caryopses 1.5-2 mm long: endosperm soli Anatomy. Leaf blades in transverse section are without a well-developed keel and with prominent ribs; furrows more than one half of the leaf thick- Volume 91, Number 1 2004 Finot et al. 13 Trisetum, Peyritschia & Sphenopholis ness; sclerenchyma present as a continuous abaxial band, adaxial girders T-shaped, marginal scleren- chyma present; adaxial epidermis has trichomes and inconspicuous bulliform cells: the abaxial epi- dermis lacks trichomes or prickles. Endemic to Chihuahua, Durango 2001). and Hidalgo. Distribution. errera- Arrieta, Mexico. 1600-3100 m. Comments. ical Trisetum species by having a terete lemma (vs. Trisetum filifolium differs from typ- lemma keeled), the lemma apex entire or bilobed (vs. 2-aristulate to 2-dentate in Trisetum). the palea enclosed by the lemma (free from the lemma in Trisetum). and by having linear lodicules (vs. 2 or 3 lobate at the apex in Trisetum). 7 a studied. MEXICO. Chihuahua: Sanchez. 12 Oct. 1910, Hitchcock 7682 (US); 27.6 mi. NW of Ro- che ү? Е оп road to Basigochi, 19 Sep. 1991, Peterson el al. 97 (US). Hidalgo: Zacualtiplán. trailside, slopes & ravines by Río P Е between Zacualtiplán & Olotla on road to Metztitlán, 3 Jul y 1947, Moore 3250 (US). KEY TO VARIETIES OF TRISETUM FILIFOLIUM the mucro 0.2-5 mm long, 3a. Т. ‘Allin v var. filifolium . Lemmas ташы. the awn 3— late and twisted, borne on бе иррег third of the lemma, i.e.. inserted е the apex (—— T. filifolium var. aristatum la. Lemmas mucronate, inserted apically 1 long, genicu- — . Trisetum filifolium var. aristatum Scribn. ex Beal. Grass. М. Amer. 2: 375. 1896. TYPE: Mexico. Chihuahua: cool slopes of the Sierra Madre. alt. 9000 ft.. 7 Oct. 1887, C. G. Pringle 1430 (holotype. US-868411!: isotypes, MO- 3727972 not seen, NY not seen). دن — = Distribution. Endemic to Mexico in the Sierra Madre Occidental in Chihuahua and Durango and Sierra Madre Oriental in Coahuila. México, and Hi- dalgo (Hernández-Torres & Koch, 1988; Finot, 2003b). Comments. ‘Though Beal (1896) suggested that variety aristatum could be a different species, the overall morphology of the plant and the structure of the spikelets conform with the general descrip- tion of Trisetum filifolium. MEXICO. Durango: 3.2 n Specimens sl udied. of 2003. Ped. ea La Pefia on road toward La Puerta, 15 Sep. et al. 17785 (CHDIR. US). Puerto Ing. н Diaz. km 230-235 on hwy. between Zimapán & Jacala, 7 July 1948, Moore & Wood. Hs 3765 (US). México: 28. 3 mi. NE of Temascaltepec on Mex towards Toluca, 7 Oct. 1991, Peterson & locals 11084 (US). 4. Trisetum irazuense (Kuntze) Hitche.. Proc. Basionym: Ca- PL Biol. Soc. Wash. 40: 82. 1927. * irazuensis Kuntze, Revis. Gen. 2: 763. 1891. TYPE: Costa Rica. Volcán lrazü, 3000 m. a 24 June 1874. C. E. О. Kuntze 2334 (holotype, NY-346300 ех Herb. Kuntze!). Trisetum 5 Hitche., Contr. U.S. Natl. Herb. 24: 358. 1927. TYPE: Colombia. Cauca: collected below Pitayó, Rio Palo Basin, Tierra Adentro, 2400 m. Feb. 1906. H. Pittier 1435 (holotype, US-53 1631). Trisetum scabrivalve т Revista Colegio Nac. Vicente а afuerte 11: 8 t, 88 (Reprint 32, 36), 1930. TYPE: ador. Crece en los pajonales del Pichincha, Pa- ai (Pifo), Chimborazo y El Altar (type not lo- cate Trisetum iE. Fourn., Mexic. Pl. 2: 108. 1886. hom. non Trisetum gracile (Moris) Boiss. Trisetum fournieranum Hitche., Contr. U.S. Natl. Herb. 17: 326. 1913. TYPE: “San Luis de Potosí,” M. Virlet d'Aoust 1382 (lectotype. ا‎ here, P!; isotype. US-72697 1b fragment ex P!). illeg.. Perennial, culms 75-100 cm tall, glabrous, up to 2 mm diam. on the lower inter- caespitose; nodes; nodes 3. glabrous. Leaf sheaths shorter or slightly longer than the internodes, glabrous or pu- bescent; basal sheaths pubescent; ligule 2—4 mm long, membranous, pubescent to densely pubes- cent, the apex truncate, dentate or ciliate: blades 20-30 ст X 2.5-6 mm, flat, glabrous or pube scent: upper blades 5-10 cm long. Panicles (7—)13-25 X (1-)2-5 em wide. green and purple; rachis glabrous to sparsely pu- lax, narrow, vellow-green to deep bescent; the branches appressed and ascending. Spikelets 4—7(-9) mm long. 2- or 3-flowered: ped- icels 2-7 mm long. glabrous to sparsely pubescent, sometimes scabrous; rachilla 1—1.7 mm long, cov- ered with stiff trichomes, the trichomes 0.5-1 mm long: glumes very unequal, shorter than the spike- et; the keel scabrous on the upper half, the mar- gins hyaline; lower glume 2.5—5 mm long, narrow, linear, l-nerved, acute at the apex. half as long as the spikelet; upper glume 3.5-6.5 mm long. lan- ceolate to ovate-lanceolate, abruptly attenuate, 3- nerved, % as long as the spikelet: lemmas 4.3—6 mm long, 5-nerved, the nerves inconspicuous to- ward the base, strongly scabrous on the upper half, green and purplish toward the apex, sometimes with short trichomes toward the base: apex awned, hyaline, biaristate or toothed, the apical awns ca. 0.5 mm long, conspicuous; awn (2.5-)5.5-8.5 mm long. borne on the upper Y or 4, straight or twisted and geniculate: callus obtuse, with short trichomes, the trichomes ca. 0.2-0.3 mm long: paleas 3.2—1.6 mm long, slightly shorter than the lemma, hyaline. 2-keeled, the keels scabrous on the upper half, 2- dentate to 2-awned at the apex: lodicules ca. 0.7 Annals of t Missouri Botanical Garden mm long, bilobed at the apex, the lobes acute; an- thers (0.8—)1.2—1.8 mm long: ovary glabrous. Cary- opses 2.2—3 mm long: endosperm soft. Chromosome numbers. 2n dez-Torres & Koch, 1988). Anatomy. Leaf blades are flat in transverse sec- 28, 42 (Hernán- tion, with rounded adaxial ribs: furrows less than one half of the leaf thickness; first-order vascular bundles with adaxial and abaxial girders; second- third- order vascular bundles with small sclerenchyma order vascular bundles with adaxial girder: strands; bulliform cells in fan-shaped groups at the base of the furrows; abaxial epidermis with fusiform intercostal long cells with straight side walls; sto- mata not seen; prickles rare; trichomes present: costal cork cells and silica bodies in pairs or short rows. Distribution. This species has been reported for Mexico (Chiapas, Distrito Federal, Durango, Hi- dalgo, México, Oaxaca. Puebla, San Luis Potosí. and Veracruz), Costa Rica, Panama, Honduras, Co- lombia, Venezuela, Ecuador, and Peru; 2700-3600 m (Pohl, 1980: Hernández-Torres & Koch. 1988: Davidse et al., 1994; Finot, 2003b). A lectotype (M. Virlet d'Aoust 1382) P was here chosen to stabilize the name since there was another syntype (Ё M. Liebmann 613) cited in the original protologue by Fournier (1886). A specimen of Liebmann 613 was not found at P. Comments. Specimens studied. COSTA RICA. San a eri e upper slopes of volcán Irazú, 24 Fel 11 5 66035 (US); San José, S-SW of Cerro М Ыш 10 De oes” Weston 3673 (CR). GUATE MAL ie ne Типи ná, Sierra de los Cue ni imatanes, 6 July 48276 (US ae _MEXICO. à 8. C. 26 971 ). Méx g 1910. chek 1448 (P). 5974 (US): 895, no lh ctor € 5 Chiriquí ups ‘ano, 29 Sep. (US), 29-30 Sep. 1911, Hitchcock Ey (S). PERU. ra: 5 i1 E of 5 аА on road toward ' e Mar. 2000, Peterson & Re "fulio- Rodriguez 15135 (US). Feb. 55. La pen fir woods, Sierra (P). PANAM 911. Hitchcock аг?! as Cruces, 5 Oc — Piu- 3l Trisetum pringlei (Scribn. ex Beal) Hitehe.. Proc. 82 Biol. Soc. Wash. 40: 1927. Ba- roba diu pringlei ar sionym: ex Beal, Grasses N. 561. . TYPE: Mexico. Oaxaca: summit of Sierra s San Fe- lipe, 10.000 ft., 4 Aug. 1894, C. G. Pringle 4765 (holotype, US-822340!; isotypes, CM not seen, ENCB not seen, MO not seen, Pl. US- 749287!, US-251958!). Perennial, e 118 culms 2 brous; nodes (1)2 or 3 25-80 cm tall. gla- . glabrous. Leaf sheaths gla- brous or pubescent, sometimes only pubescent at the apex, shorter or longer than the internodes, sometimes one side extended above summit as a sheath auricle: ligule 0.5-1.5 mm long, the apex truncate to ovate, minutely denticulate and ciliate: 4-9(-12) cm X 2-5 mm. flat. subinvolute, the lower blades glabrous or pubes- blades sometimes cent, ciliate on the margins, the upper blades gla- Panicles 4—17 —12 cm, open, lax, usu- ally purplish; branches up to 8 em long, verticillate, capillary and flexuous: rachis glabrous to somewhat scabrous. Spikelets 3.5-6.5 mm long, 1- or 2(3)- flowered, pedicels up to 4 mm long, brous. capillary, g a- brous, purple: rachilla 1-1.5 mm long, with tri- chomes 0.5-1 mm long. distributed toward the apex: glumes unequal, shorter than the spikelet, acute, the keel smooth or scabrous on the upper half: first glume 1.4-2.6 mm long. linear-lanceo- late, 1- nerved; second glume 2-4.5 mm long. lanceolate, 3-nerved: shorter and narrower than the second. lemmas 2.6-4 mm long, strongly sca- rous, sometimes covered by short trichomes, pur- ple. terete, 5-nerved, the nerves conspicuous to- ward the apex: margins hyaline; apex truncate, entire or mucronate; subapical mucro when present less than 1 mm long: paleas 2.2-2.9 mm long. shorter than the lemma, hyaline, 2-nerved, some- times purple-tinged, the keels scabrous to ciliate- callus obtuse, with tri- chomes 0.1-0.5 mm long: lodicules 0.4-0.7 mm long. linear, not lobed, the apex obtuse, ciliate: apex rounded or acute; anthers 0.6-1.2 mm long: ovary glabrous. Caryopses I. EI.) mm long. ca. 0.4 mm wide: endosperm solid. scabrous on the upper half; Anatomy and micromorphology. Leaf ligule apex has only papillate cells and the epidermis is composed of more ~ г less rectangular long cells with straight side walls; hooks and prickle hairs present; stomata Leaf blades in transverse section are without a keel and have well- developed ribs: furrows more than one half of the leaf thickness. bulliform cells inconspicuous; vas- were not seen. cular bundles with both adaxial and abaxial girders alternate with free bundles without associated scle- renchyma, adaxial girder T-shaped, abaxial girders I-shaped, marginal tri- chomes on both adaxial and abaxial surfaces. sto- sclerenchyma_ present: mata present on the adaxial epidermis; abaxial epi- dermis composed of long cells with undulate side walls: costal cork cells-silica bodies present, tri- chomes present, prickles not seen. Distribution. la), Guatemala, Costa Rica, and Panama: 3500 m (Hernández-Torres & Koch. 2003b). Mexico (Chiapas, Oaxaca, Pueb- 1500— 1988; Finot, Volume 91, Number 1 2004 Finot et al. 15 1 950 Peyritschia & Sphenopholis Comments. The number of florets in Trisetum pringlei varies from one to three per spikelet. One- flowered spikelets are infrequent in Trisetum, and are found in some individuals of T. filifolium, T. pringlei, and in the South American 7 ambiguum Rúgolo & Nicora (Rúgolo de Agrasar & Nicora. 1988). On the basis of morphological, anatomical, and embryological characteristics, Hernández-Torres and Koch (1988) concluded that this species be- longs in Trisetum, as was proposed by Hitchcock (1927). Trisetum pringlei differs from typical spe- cies of Trisetum by having lemma apices that are entire and mucronate, and linear lodicules with en- tire (not lobed) apices. Further studies are needed to clarify the generic assignment of T. pringlei (see Appendix 1). COSTA RICA. Cartago: along the Pan dip Iwy.. Cerro de la Muerte, 8 Aug. 1972, Taylor 11750 (US): Crater volcán lrazü. a Oct. Anderson 1339 (US-2042177): Cerro de la Specimens studied. American Hwy., 5 km above Mill: Isville, UM ra de Ta- m nca, 22 July 1949, кезе des Iltis 483 (P). San José: ordillera de Talamanca, o de La Muerte, Asunción summit, 13 July 1968, d € "Davidse 10694 ( CR). GU. A- TEMALA. Huehuetenango: b b Ў bluff. summit of Sierra de los Cuchumatanes, 6 Aug. 1942, Steyermark 50224 Chiu Jolán, mounts above Totonk 19 (US). Totonicapán: region of ps i ау to Des- Standley ). PANAMA C hiriquí: C ШТ Mg 'ano, 2t E т E р: is l. Hite ба 228 (US). eo Jaxaca: bois of Sierra de San Felipe. 4 i 89: кл. E : vicinity of Cerro San Felipa, vi sli 1108 (US consuelo, 2: И. 6. Trisetum barbatipaleum (Hultén ех Veld- kamp) Finot, Contr. U.S. Natl. Herb. 48: 661. 2003. Basionym: Trisetum spicatum var. bar- batipaleum Hultén ex Veldkamp, Gard. Bull. Singapore 30(1): 135. 1983. Trisetum spicatum subsp. tolucense var. barbatipaleum Hultén. Svensk Bot. Tidskr. 53: 223. 1959, nom. inval. TYPE: Mexico. Hidalgo: Trinidad Iron Works, 1770 m, 21 Aug. 1905, C. G. Pringle 10052 (lectotype. designated by Veldkamp, in Veld- kamp & Van der Have (1983: 135). S not seen; isotypes, C not seen, L not seen, PL SE. 162083!). Perennial, caespitose; culms ca. 40 em tall, pu- bescent; nodes pubescent. Leaf sheaths shorter than the internodes, pubescent; blades ca. 7 cm X 1-2 mm. flat, pubescent abaxially, glabrous ada- xially. Panicles 8-10 X 0.5-1 em, spiciform, in- terrupted; rachis pubescent: pedicels pubescent, Spikelets 5-5.2 mm long, 2-flowered: rachilla ca. | mm long. pilose; glumes unequal, shorter than the scabrous on the keel: first spikelet, somewhat glume 3-3.2 mm long, shorter and narrower than the second glume, linear-lanceolate to lanceolate, slightly longer than half of the spikelet, I-nerved; second glume ca. 4 mm long, ovate-lanceolate to ovate, 25 as long as the spikelet, 3-nerved: lemmas 3.5-5 mm long, pubescent, 2-aristulate at the apex. awned on the upper third; awn ca. 6 mm long. curved, divaricate, not geniculate or twisted, ciliate near the base and scabrous above: callus with short trichomes, the trichomes ca. 2 mm long: paleas shorter than the lemma, ciliate toward the apex. Lodicules ca. 1 mm long, bilobed at the apex; ovary glabrous. Caryopses not seen. Distribution. Endemic to Mexico where the species is known only from the type locality. Comments. Trisetum barbatipaleum differs from T. spicatum in having conspicuously pubescent lemmas. It is closely related to 7: rosei and differs from the latter by having a narrower panicle, glumes that are shorter than the spikelets and sca- brous along the keel, and a first glume that is nar- row and linear-lanceolate. Trisetum rosei has ovate glumes as long as the spikelet, a wider and more densely flowered panicle, glumes that are conspic- uously ciliate on the keel, and a lemma with a plu- mose awn. 1. Trisetum ligulatum Finot & Zuloaga, sp. nov. TYPE: Mexico. Veracruz: La Perla, N side of Pico de Orizaba, above the Piedra Grande mountaineering shelter, 19503“ N. 97716'W, 4200 m. 21 Sep. 1986, M. Nee 33192 (holo- type. SI!; isotype, CONC!). Figure 1. Planta rhizomatosa, 13-30 em alta; culmi erecti, pilosi vaginae pilosae; ligula 3—4 mm longa, Malina, truncata, dorso pilosa: laminae planae 5-20 ст X 1-2.5 mm), pi- losae; panicula spiciformis; spicula pa 5 mm le 2- 2 3-flora; glumae subaequales. inferiores (1)3-nervatae, su- nata scabra, 5-nervata, ad e biaristato; arista 2-3.5 reve pilosus; palea paulo brevior quam — ian (3)5-nervatae; lemi — ~ = ‹ 1 superiorem aristata, apice mm быз. callus b lemma: lodiculae ad apicem 3-lobatae; ovarium glabrum. Perennial, with short rhizomes; culms 13-30 cm tall, erect. pilose below the panicle, the trichomes antrorse above then retrorse below: nodes 1 to 3. — glabrous. Leaf sheaths longer than the internodes, pubescent; ligules 3-4 mm long. dorsally pilose, truncate: blades 5-20 em X 1-2.5 duplicate, pubescent. Panicles 5-9 X 0.5-1.2 cm. 5 mm. flat or con- spiciform, green or purple; rachis densely pubes- cent; spikelets 5-6.5 mm long. 2- or 3-flowered: rachilla ca. 1 mm long, sparsely pubescent, the tri- chomes 0.5 mm long; glumes subequal, nearly as long as the spikelet, acute, the keel scabrous on the upper half, green or purple: first glume 3.8—6 16 Annals of the Missouri Botanical Garden Mas, EAN SS Dh — = ee a B Е 3 N o A gure en id و‎ (Nee 353192). —А. Habit. —B. Sheath, ligule, and portion of the blade. C. Spikelet. . Floret. —F. Palea, dorsal view. —G. Palea. lodicules, and stamens, ventral view. m Pistil. mm long, as long as or shorter than the first floret, nearly as long as its lemma, scabrous on the keels: (1)3-nerved: second glume 4.5-6.5 mm long. (3)5- — lodicules 3-lobed at the apex: anthers ca. 1.5 mm; nerved; lemmas 4.3-6.5 mm long. scabrous, 5- ovary glabrous. Caryopses not seen. nerved, keeled, awned: margins hyaline toward the apex: apex biaristate: awn inserted on the upper Anatomy. Leaf blades in transverse section are third, 2-3.5 mm long. curved. scabrous: callus without a keel and have well-developed ribs with short pilose, the trichomes 0.5 mm long: palea furrows more than half the leaf thickness: bulliform Volume 91, Number 1 2004 Finot et al. Trisetum, Peyritschia & Sphenopholis cells are inconspicuous, trichomes present only in the abaxial epidermis, continuous abaxial scleren- chyma band absent, vascular bundles of first order without girders, bearing small strands of both ad- axial and abaxial sclerenchyma. marginal scleren- chyma present: stomata present in both adaxial and abaxial epidermes; abaxial epidermis with rectan- gular to fusiform long intercostal cells, short costal cells solitary or in cork cell-silica body pairs. prickle hairs present. Distribution. This new species is known only from the higher diesen (3810—4200 m) of Jalis- co and Veracruz, Mexico Comments. Trisetum ligulatum resembles T. spicatum in having spiciform panicles and culms pilose below the inflorescence. It differs from 7. spicatum in having short rhizomes (vs. none), dor- sally pubescent ligules 3—4 mm long (vs. glabrous ligules ca. 1 mm long). spikelets with the first glume 3-nerved and the second glume 5-nerved (vs. first glume I-nerved and second glume 3-nerved, and with both glumes nearly as long as the spikelet (vs. shorter than the spikelet) MEXICO. Jalisco: Nevado de Colima, 2t mi. SW of Ciudad Guzmán and 14 mi. SW of paved hwy Paratype. ур road toward the Nevado, 23 Nov. 1995, P. M. Peterson R. Blackburn 13778 (US 8. Trisetum rosei Scribn. & Merr.. Contr. U.S. Natl. Herb. 8(4): 289. 1905. Trisetum rosei var. tenerum Scribn. & . Contr. U.S. Natl. Herb. 8: 289. 1905. "gs rosei Scribn. & Merr. fo. tenerum (Scribn. & Merr.) Louis-Ma- rie. Rhodora 30: 239. 1929. TYPE: Mexico. Vole: ‘ап Popoc ow 3600 m. 7-8 Aug. 1901, J. — Rose & К. : 6016 (holotype, US- 3958131: isotypes. ENCB not seen, MEXU- 4703 not seen, NY-431715 not seen). Perennial, caespitose: culms 25-50 cm tall. densely pubescent, the trichomes antrorse above then retrorse below: nodes 2. Leaf sheaths glabrous or retrorsely tomentose. shorter than the internodes; ligule 1-2.5 mm long, scabrous on the dorsal sur- face, truncate, conspicuously ciliate; blades 6-15 cm X 2-4 mm. flat, glabrous to pubescent on both sides, soft or stiff. conduplicate or the margins in- volute, the uppermost leaf blade 3—6 cm long. Pan- icles 3 * 1-1.5 cm, subspiciform, dense. + linear, acute at the apex, sometimes interrupted; rachis pubescent. Spikelets 5-7 mm long, 2(3)- flowered: pedicels up to 3 mm long: rachilla ca. 1 mm long, pubescent, the trichomes stiff, са. 0.5 mm long: glumes slightly shorter to as long as or longer than the spikelet, ciliate on the keel, the margin hyaline, the apex acute, faintly tinged with purple: first glume 3.7—6 mm long, lanceolate to ovate-lan- ceolate, usually 1- or 3-nerved; second glume 4.6— 6.5 mm long, ovate-lanceolate to ovate, 3-nerved: lemmas 4.2—6.2 mm long, pubescent, awned: apex bisetulate and hyaline, seta 0.3—0.7 mm long: awn 2.5-5 mm long, borne on the upper third. plumose. curved, not twisted or bent; callus obtuse, with tri- chomes 0.3—0.5 mm long; paleas 3—4.5 mm long. glabrous to pubescent, shorter than the lemma, 2- nerved, the keels ciliate: apex bidentate, hyaline: lodicules 0.5-1 mm long, 2-lobed at the apex, the lobes unequal, acute; anthers 1-1.5 mm long; ovary glabrous. Caryopses 2.5-2.7 mm long. ca. 0.6 mm wide; endosperm liquid or soft. Anatomy and micromorphology. Ligule apices have short stiff trichomes, and the epidermis is composed of more or less rectangular long cells with straight side walls, hooks very frequent: no Leaf blades in transverse section did not have a keel but had well- trichomes or stomata were seen. developed ribs with furrows more than one half the leaf thickness; bulliform cells inconspicuous; tri- chomes present in both adaxial and abaxial epi- dermes with a continuous abaxial sclerenchymatic band present; first- and second-order vascular bun- dles with adaxial and abaxial girders; adaxial gird- ers T- or anchor-shaped; marginal sclerenchyma present; abaxial epidermis with long cells with un- dulated side walls, intercostal hooks and stomata present, cork cells and silica bodies present in cos- tal zones; no trichomes were seen in the abaxial epiderm a México, Puebla, and Veracruz), between 2500 and 4300 m (Espejo-Serna et al., 2000). Comments. Guatemala and Mexico (Hidalgo. Trisetum rosei was considered a synonym of T. spicatum by Hernández-Torres and Koch (1988). Trisetum rosei can be easily recog- nized by its pubescent lemmas, ciliate glumes, plu- mose awned lemmas, paleas with ciliate keels (vs. lemmas glabrous, glumes scabrous, lemma awns not plumose, and scabrous palea keels in J. spi- catum), and, in cross section, an abaxial band of continuous sclerenchymatous tissue (absent in 7. spicatum). e studied. GUATEMALA. Huehuetenango: s Chuchumatanes, 6 July 1942. Steyermark m ME XICO. Hildalgo: Inter Pachuca et Min- eral del Chico, 20 Dec. 1931, зан & Hultén 3 (5): mtns. above Pachuca, 4 Aug. 1 ] J > tween Pachuca & x del ^ n 19 Rose. Painter & J. S. Rose 8686 (US). México: V Ixtaccihuatl, media falda 13 Sep. 1953. Matuda 29044 (US); 24.2 km E of Amecameca & 2.5 km ' S of Paso de Annals of the Missouri Botanical Garden 10 Oct. 1991, e Cortéz. Pe 7 & Annable 11103 ( 1958, Beaman oe (U n Popoc ate E 5-6 iub 1910. E 87 (US): don | 9 km E of San 1 16 July Puebla: № side of Popocate- 1729a (US); ca. 1 km Paso e UR Mt. Orizaba. US): Mpio. San Ni- 1, Vol ‘ап Ixtaccihuatl, 7 km n E Amecameca, 2 pee Koch 7623 1 (US); Mpio. de Tochimilco, du Cor- 7. Tlamacos, 16 km ESE de Amec n a, 18 Oct. 1976, pan 762056 (US). Veracruz: Mt. Orizaba, 15 1908. Purpus 3018 (BAF); Pico de Orizaba, N of Cueva del Muerto, 21 Sep. 1957, Beaman 1778a (US) side of mt. ca. 1959, ‘ama 2853 (US). petl, Sep. de C 1 z, 21 July 1 1910, „д — ~ Jet. 9. Trisetum spicatum (L.) K. Richt., Pl. Eur. 1: 59. 1890. Basionym: Aira spicata L., Sp. Pl. Aira subspicata L., Syst. Nat. ed. 10 2: 873. roides Koeler, Descr. 159, nom. Шер. superfl. Avena ai- Gram. 298, 1802, nom. illeg. superfl. Trisetum subspicatum (IL) P. Beauv., Ess. Agrost. 88, 1812, nom. illeg. superfl. Trisetaria airoides (Koeler) Baumg., Enum. Transsilv. Stirp. 3: 265. 1816, nom. illeg. superfl. Trisetum airoides (Koeler) P. Beauv. ex Roem. & Schult., Syst. Veg. 2: 666. 1817, nom. illeg. superfl. Koeleria subspicata (L.) Rehb., Fl. illeg. superfl. Koeleria spicata Rchb. ex Wi & Lange. Prodr. Fl. Hispan. 1: 72. 1861, nom. inval. Trisetaria spicata (L.) Paunero, Anales adrid 9: 516. 1959. TYPE: Swe- Lapland: 1732, Linnaeus s.n. (lectotype, Germ. Excurs. 30, nom. designated by Edgar € Connor, in Edgar (1998: 556). LINN-85.7!; isotype, Sh). Avena tolucensis Kunth, Nov. Gen. Sp. 1: 148. dH Tri- setum tolucense (Kunth) Kunth, Havin: Gram. 1: 101 t. 60. 1829. Trisetum spicatum acu (Kunth) Hultén, Sve ШЕ 1 Bot. Tidskr. ы PE: Mexico: "crescit in apricis, op ee ur- bem mexicanam Тос а, alt. 1380 he Нит- ae & Bonpland s.n. (holotype, P!; 5 8 BAA- 9 fragment ex Kl. BAA-3420 fragment ex Р!). S n nivosum E. dapi кт PI. 2: 107. 1886. Tri- setum spicatum var. ni n (E. Fourn.) bs MO ) 1928 1 10201 TYPE: Mexico. Ne- vado de Toluca, Sep. 1865, Hahn s.n. (lectotype. designated by juan аш 688), Pl: isotype, US- fragment ex Р-Еош 2 ense . 1959 Perennial, caespitose; culms 9—50 cm tall, erect, densely pubescent below the inflorescence, the tri- chomes antrorse above, retrorse below: nodes 1 or 2. Leaf sheaths 1—3(—6) cm long, glabrous: ligule 0.5-1 mm long, minutely denticulate; blades 1—4 em X 121.5 mm, flat, conduplicate toward the apex, glabrous to pubescent, sometimes scabrous or ciliate on the margins, the upper blade 1—5 em 2.5-1(-10) х 0.5-1.3(-2) em, spi- long. Panicles 2 spicatum var. ciform, green to purple, shining, usually interrupted on the lower portions; rachis pubescent. Spikelets 1.5-5.5 mm long. (1)2-flowered: pedicels pubes- cent; rachilla ca. 1 mm long with trichomes 0.5- mm long: glumes subequal, wide, shorter than the spikelet, or the first glume somewhat shorter and narrower than the second glume, scabrous or more rarely ciliate on the keel; first glume 3.7—4.8 mm long, lanceolate to ovate-lanceolate, l-nerved; sec- -— ond glume 4.5-5.5 mm long, ovate, 3-nerved; lem- mas 3.8—5 mm long, glabrous, purplish toward the base, awned; margins hyaline; apex 2-aristate; awn 3.5-5 mm long. borne on the upper third or upper fourth, geniculate to curved, sometimes slightly twisted, scabrous, purplish; callus obtuse with tri- chomes 0.3 mm long: paleas 3—4 mm long, free from the lemma, shorter to slightly longer than the lemma, hyaline, 2-nerved, the keels scabrous; lod- icules 0.6 mm long, hyaline, 2-lobed at the apex: ovary glabrous. Caryopses 2-2.8 mm long, ca. 0.6 mm wide; endosperm liquid. Anatomy and micromorphology. Ligule apices have long cells and papillate cells with trichomes absent; epidermis with rectangular long cells with straight side walls, without trichomes, hooks, or sto- mata. Leaf blades in transverse section have prom- inent ribs with furrows deeper than one half the leaf thickness: adaxial and abaxial girders present: marginal sclerenchyma present; adaxial epidermis with inconspicuous bulliform cells; abaxial epider- mis without trichomes and only few prickles pre- sent Distribution. Cosmopolitan and widespread. Hultén (1959) and Clebsch (1960) gave a detailed account of the distribution of Trisetum spicatum. In Mexico and Central America T. spicatum is usually found between 1900 and 4500 m Comments. Trisetum spicatum has been consid- ered a complex, with many subspecies or varieties (Louis-Marie, 1928—1929; Hultén, 1959). Louis- Marie. recognized 14 varieties and Hultén (1959) divided J. spicatum into 22 тере laxa, in- cluding 14 subspecies and 8 varieties. In a phe- netic study, Randall and Hilu (1986) concluded that the morphological variation is extreme and - -= does not support the recognition of infraspecific taxa. In this treatment we interpret 7! spicatum in a stricter sense, revalidating T. rosei and raising T. barbatipaleum to specific status. However, T. spicatum remains an extremely vari- able taxon with a large number of synonyms. In this revision we have included only those synonyms that occur in the study area, in addition to the basionym and its nomenclatural synonyms. Volume 91, Number 1 2004 Finot et al. 19 Trisetum, Peyritschia & Sphenopholis DOMINICAN REPUBLIC. Distri- to Nacional: Santo Domingo, Tierra de Oco Azua. San José de Ocoa. Cuchilla del Pino n Mar. 1929, Ekman 11214 (S): Cordillera Central, Prov. i a е Pico del Valle Nuevo, 15 Oct. 1929, Ekman 13759 (S). oie Cordillera de Baharuco, LM чта de Los о 29 Е g. 1926, Ekman 6803 (S). MEXI- CO. Durango: in montibus prope aguae stagni seu кы Aug. 1857, 0 ا‎ 57 (Р). dales o: Sierra de Pachuca, 21 July 1901, Pringle 9603 5 Sep. 1958, Hernández 25 Sep. Specimens studied. Prov. ii Convalli ma ani Us). ud ө: Nevado de Toluca, 5 X 1019 т Pringle 4303 (S, 5) 19( : Nevado de Toluca, en el « 70 05 Nevado de Toluca. crater, . Diguet s.n. Mt. Popoc ано Hitchcock 59% с, Valle de V o, falda de Ixtaccihuatl, | May Matuda ыш (US); Nevado de Toluca. \ lb Balls WE 5 (US): Volcán Toluca, 9 Sep. 1893, Nel- son 17 (US : Таас C ‘ihuatl, above timberline. Oct. sco: Nevado de Colima. 26 Aug. . Orizaba. between cakes and snow line, Mucha 6258 (US 5): Mt. alpine region, Nov. 1906, Rose 1270 (US). A N хы со > ® — a = = e 8 a E E № e ~ NX c Orizaba, 10. Trisetum durangense Finot & P. M. Peter- TYPE: - Me хїсо. Madre Occidental, ' of Durango on Hwy. 40, 2400 m, 29 Sep. 1988, P. M. Peter- son & C. R. Annable 6034 (holotype, US!). Fig- ure : son. sp. nov. ‚ Durango: Sierra Hs Sigs pilosae, inte odia paulo breviores vel aequale — anta perennis: culmi glabri vel scabri sub casi ligula 3-67) mn Аарне alae, Ше, 15-50 cm X supra scabrae vel эра rse рне paniculae 15 a 20 cm. laxae: rhachis scabra: spiculae 7-9 т 2(3)- bu ume bee "n quam мйне ча contigua gluma боз acuta, lacerata; laminae con- n longae. minore 5; superior brevior angustior- aristulata: callus pilosus; rhachilla pilosa. 2 mm longa: palea 5.8-7 mm longa: lodiculae 0.7—1.1 mm longae: sta mina 3: ovarium ad apicem breve pilosum. „Perennial, caespitose, short rhizomatous: culms —110 cm tall. ы the panicle: nodes 3. pubescent. Leaf sheaths glabrous or retrorsely scabrous be- 11-23 cm long. pilose, as long as or shorter than the internodes, with the membranous margin on one side sometimes projecting upward as long as a sheath-auricle: ligule 3—6(—7) mm long. acute, lac- erated above: blades 15—50 em X 1—4 mm, narrow, flat to conduplicate, rigid. strongly scabrous on the abaxial surface. scabrous and sparsely pubescent on the adaxial surface, ciliate on the margins. Pan- icles 15-22 X 5-20 cm. lax, drooping, open, py- ramidal: branches appressed or ascending to spreading 0 to 75° from the culm axis, branches in 4 or 5 verticillate whorls per panicle, each whorl composed by 4 to 6 branches, apical branches 3— 7 em long. the lower branches 8—14 cm long: ra- chis. branches and pedicels strongly scabrous, cap- illary, bearing spikelets on the distal ends. Spikelets 7-9 mm long. 2(or 3)-flowered: glumes equal to subequal, similar in shape, as long as the spikelet or slightly shorter than the spikelet (first): first glume 5-6 mm long, lanceolate, slightly short- er and narrower than the second glume, 3-nerved. scabrous on the keel, apex acute: second glume 6— 9 mm long. lanceolate, 3-nerved, apex acute: lem- mas 7-8 mm long. awned near the middle. pilose especially below the awn insertion, 5-nerved, the nerves conspicuous toward the apex but vanishing toward the base of the lemma: margins of the lem- ma flat; apex 2-lobed, the lobes acute with the in- termediate and marginal nerves prolonged into 4 short apical awns; awn 7—10 mm long. geniculate and twisted, golden brown, exserted: callus pubes- cent, the trichomes ca. 1 mm long; rachilla ca. 2 mm long, pubescent; paleas 5.8-7 mm long. hya- line, 2-nerved, the nerves ciliate; lodicules 0.7-1. mm long. ciliate at the apex; anthers 2-3 mm long. curved trichomes; ovary ca. 1 mm long. covered with short, curved and shining trichomes near the apex. Caryopses 2-2.7 ca. 0.7 mm: endosperm soft. Anatomy and micromorphology. ligule apex composed of trichomes and some papillose cells: ligule epidermis composed of long cells with straight walls; prickles absent on the dorsal surface. only present at margins; macrohairs and stomata absent. Transversal section of the leaf blade with conspicuous adaxial ribs: vascular bundles with ad- axial girders T-shaped; abaxial girders united form- ing a continuous band of abaxial sclerenchyma; ab- axial epidermis with macrohairs and abundant prickles; epidermal long cells with undulate lateral walls; short cells present in costal and intercostal zones; stomata absent in abaxial epidermis. Distribution. Known only from the state of Du- rango, Trisetum durangense occurs in open pine for- ests between 2400 and 2600 m. Comments. Trisetum durangense is distin- guished by its pilose sheaths; pubescent nodes: strongly scabrous blades: panicle with rachis, branches, and pedicels strongly scabrous: long, pi- lose lemmas; and an ovary with short, curved, and shining trichomes at the apex. The presence of these apical trichomes on the ovary is found in sev- eral taxa, including T. cernuum Trin. and J. canes- cens Buckley from North America, and T. hirtiflo- rum Hack., T. caudulatum Trin. var. correae Nicora, and T. longiglume Hack. var. longiglume from South America. Trisetum durangense is the only species in subgenus Deschampsioidea with pubes- cent lemmas and with an ovary with curved and 20 Annals of the Missouri Botanical Garden \ \ | Ji NA ril M UN 11, M NN j M | | WN \' \ N \ | | | Y | | | — SSS == — Innable 6034). —A. Habit. —B. Inflorescence. —C. Sheath, ligule, . First glume, dorsal view. —G. Second glume, dorsal view. 2). —L. Stamens (3). — Figure 2. Trisetum durangense (Peterson & + and portion of the blade. —D. Spikelet. —E. Floret. —F. —H. Lemma, dorsal view. —I. Palea, dorsal view. —J. Palea, ventral view. К. Lodicules ( M. Pistil. —N. Caryopses (2). Volume 91, Number 1 2004 Finot et al. 21 1 Peyritschia & Sphenopholis shining trichomes near the apex. Trisetum duran- gense appears closely related to T viride and can be easily distinguished from the latter by possess- ing pilose lemmas (glabrous in T. viride) and pu- bescent ovaries (glabrous in T. viride). Paratypes. MEXICO. ри Mpio. de Durango. Parque El Tecuán, 58 km W of Durango, 20 Sep. 1984 F. Casillas, R. Flores & Е Ruiz 9 (CHDIR); 56 km W of Mie on Hwy. 40 toward Mazatlán, 23%54'45.1 105501540. O W. 6 Oct. 2002, M. Peterson & L. Brothers 16964 (CONC, US); ca. 28 mi. N of Unidos aoe ceremos on road toward San Miguel de Cruce 24°17'49.8"N, 105?29'32.2"W, 7 Oct. 2002, P. M. Peter- son & L. E. Brothers 16985 (CONC, US). 11. Trisetum martha-gonzaleziae P. M. Peter- son & Finot, sp. nov. TYPE: Mexico. Durango: El Mezquital, 25 km de Los Charcos por ca- mino a La Guajolota, 2110 m, 14 Mar. 1985. M. González-Elizondo et al. 1645 (holotype. CHDIR). Figure 3. Planta perennis: culmi glabri, striati, 3-nodi, 75—100 cm alti; vaginae glabrae, striatae, quam internodia brevio- ; ligula 0.5-1 mm, ovata: laminae planae vel 77 ae, ages cn panicula contracta, acuta, 10-20 X 1.5-2(— 4) em: spiculae biflorae (triflorae). 7.5-9 mm pen glumae « mm; lemmata glabra ad apicem bidentata: callus pilosus: ovarium pilosum. quales, 7.5-9 X 0.8-1.1 Perennial: culms 75-100 cm tall, glabrous. stri- ate, nodes 3. glabrous, dark. Leaf sheaths 3-10 cm long, glabrous, striate, shorter than the internodes, ciliate on the margins near the summit; ligule 0.5 ovate, long, vegetative culms, mm slightly cleft at the apex, minutely denticulate-cil- longer in iate at the apex, glabrous on the dorsal surface: blades 10-30 em X 2-3.5 mm, stiff, scabrous on both sides. 1.5-2(-4) tinged with gold and purple, the branches semi- flat or convolute. Panicles 10—20 X em, contracted, subspiciform, acute, verticillate, each whorl composed of 5 or 6 branch- es, the lowest branch up to 5 cm long; rachis gla- Spikelets 7.5-9 mm long. 2(3)-flowered: pedicels glabrous; rachilla 1.4-1.5 mm long, densely pubescent, the trichomes ca. 1.5 mm long: glumes 7.5-9 mm long. 0.8-1.1 mm wide, equal, longer than the spikelet, weakly purple at the base, yellow at the apex, the keel slightly scabrous, the brous. = apex acute; lemmas са. 5 mm long, glabrous, 5- nerved, rounded in cross section, bidentate at the apex. the intermediate nerves reaching the apex or prolonged beyond the apex as very short hyaline selae; awn ca. 5 mm long. inserted (1.5 mm from the base) near the middle or lower third, strongly twisted, geniculate, as long as the lemma; callus obtuse, pilose, the trichomes 1-1.2 mm long: palea ca. 5 mm long. about as long as or longer than the lemma, 2-nerved, the nerves well apart, tinged with purple and slightly scabrous: lodicules ca. 0.5 mm long. bilobed at the apex: ovary pilose. Distribution. Endemic to Mexico (Chihuahua and Durango) between 2110 and 2450 m. Parques. MEXICO. Chihuahua: 53 mi. W of Balle- za and 5 . E of Guachochi, 19 Sep. 1991, Peterson et al. 10784 (US). Durango: Mpio. El Mezquital. a 5 km del a del camino de Charcos a El Mezquital, 19 Mar. ‚ O. García 82 (CHDIR); ге а ие за терен, de Santa María Ocotän. Nov. . M. Gon zález-Elizondo s.n. (CHDIR) Etymology. This species honors the Mexican botanist Martha González-Elizondo. collector of the type specimen. 12. Trisetum palmeri Hitchc., Contr. U.S. Natl. Herb. 17: 325. 1913. TYPE: Mexico. Durango: Otinapa. 25 July-5 Aug. 1906, E. O. Palmer 342 (holotype, US-571365!; EHI. MO-3056871!). isotypes, Perennial, culms 60-100 cm tall, glabrous. Leaf sheaths glabrous or scabrous; ligule caespitose; 3-6 mm long, truncate, dentate, conspicuously cil- iate at the apex, glabrous on the dorsal surface: leaf blades 10-20 ст X 1-3(25) mm, stiff, scabrous, sometimes glaucous. Panicles 10— 20 X 2-6 cm, the branches 4 to 6, verticillate, the branches in distant involute or flat. lax, open, somewhat narrow, whorls: rachis and pedicels glabrous. Spikelets 5— 6.5 mm long, 2- or 3-flowered: rachilla ca. 2 mm the trichomes 0.5-1(-1.4) mm long: glumes subequal, both glumes shorter than the spikelet, the margin hya- line. the keel smooth or only slightly scabrous; apex acute; 3.5-4.5 mm long, slightly shorter and narrower than the second; sec- ong, 3-nerved; lemmas 4—5.3 pubescent, long, linear-lanceolate. first. glume | -nerved, ond glume 4—: mm long, somewhat fragile, translucent, terete, sca- brous toward the apex, awned, the awn borne at the middle or upper third, 5-nerved, the nerves con- spicuous below and vanishing toward the apex, or more rarely prolonged in 4 very short awns; apex 2-lobed, the lobes obtuse, sometimes erose; awn 5— 7 mm long, geniculate and twisted, exserted: callus with trichomes, the trichomes 0.5-1 less; paleas 4—5.5 mm long, equal to slightly longer than the lemma, free: anthers 2-2.5 mm long: ovary glabrous. Caryopses not seen. mm long or Anatomy and micromorphology. ligule apices have short stiff trichomes and some papillate cells. epidermis with long cells rectangular to fusiform walls, tri- with straight side hooks present: no 22 Annals of the Missouri Botanical Garden if °з D A H » re = E uini n gonzaleziae (Gonzalez “ а 1645). —A. Habit. —B. Sheath, Sio and portion of the bla de. — C. Spikelet . Florets (2). —E. Floret. —F. Palea with rachilla, dorsal view. —G. Palea, lodicules, pistil, and stamens, 1 view. ede Pistil. Volume 91, Number 1 2004 Finot et al. 23 Trisetum, Peyritschia & Sphenopholis chomes or stomata were observed. Leaf blades in transverse section are without a well-developed keel but have well-developed ribs: furrows more than half the leaf thickness, bulliform cells in as- semblages of 4 to 9 in fan-shaped groups at the bottom of the furrows; trichomes present on the ad- axial epidermis only: continuous abaxial scleren- chyma band present; first- and second-order vas- cular bundles with adaxial and abaxial girders, the adaxial ones T-shaped, marginal bundles only with adaxial girder; stomata present on adaxial epider- mis only, marginal sclerenchyma present; epider- mis showed costal/intercostal differentiation, hooks both in costal and intercostal zones, cork cells and silica bodies in pairs; no stomata or trichomes were 1. Distribution. Endemic Mexico (Chihuahua, Coahuila, Durango, Jalisco, Nuevo León, Sinaloa, between 2700 and 3400 m (Espejo- 2000; Finot, 2003b). Hitchcock (1913) pointed out that Trisetum palmeri is closer to Deschampsia than to — and Sonora Serna et al., Comments. Trisetum; however, he described the species under Trisetum on the basis of the glumes shorter than the 3 florets. Trisetum palmeri was synonymized with T. viride by Hernández-Torres and Koch (1988), from which it differs in having more rigid leaves and a longer ligule (3-6 mm long). Trisetum viride has a short ligule (0.5—1(—2) mm long). MEXICO. Coahuila: 55.3 km SE Specimens studied. of Saltillo and 16.6 km SE of Jame on road to Sierra La Viga. 26 Sep. 1990, Peterson et al. 10065 (US). Durango: Sierra Madre poco 2 mi. N of Hwy. | „ Peterson el al 13424 (U СИН. huana and 4.7 m. Mesa ы Clo ria, 11 Sep. 2003, « E Sánchez-Alvarado 17743 (€ ae US). uevo León: Sierra Madre Oriental, 11.4 mi. W of Die- де; ‘ho de Marzo up road toward Cerro At 18 Oct. 5). 995. Peterson & Knowles 13.336 Peterson 13. Trisetum pacas Swallen, Phytologia A(T): 424. TYPE: Guatemala. Quezal- tenango: 9 Santo Tomas, 22 Jan. 1940, J. A. Steyermark 34824 (holotype, F- 1048257 not seen; isotype, US-2230478 ex Fh. Perennial: culms 35-70 cm tall, glabrous. Leaf sheaths glabrous. with the margin extended up as a sheath-auricle 3 mm long. as long as or longer than the ligule; ligule 2-3 mm long, hyaline, trun- cate, the apex denticulate, glabrous on the dorsal surface; blades 1-2 mm wide, flat, glabrous. Pani- cles 8-15 X 0.6-1 em, spiciform, narrow, densely flowered, interrupted, green; branches 1—3 cm long, closely appressed and ascending: rachis glabrous to slightly scabrous. Spikelets 6.5-7 mm long, 2- flowered; glumes subequal, ovate-lanceolate, acute at the apex, green, the margins hyaline, the keel slightly scabrous, both glumes slightly shorter than the spikelet; first glume 5-6 mm long, l- to 3- nerved; second glume ca. 6 mm long, 3-nerved: lemmas ca. 6 mm long, scabrous near the apex, 5- nerved, awned from the middle or lower third, the margins hyaline and involute below, the apex hy- aline and deeply bidentate, with 4 hyaline setae formed by the prolongation of the intermediate and marginal nerves beyond the apex: awn 10—12 mm long. geniculate and twisted; callus obtuse with tri- chomes ca. 1 mm long; paleas 3—4 mm long, shorter than the lemma and enclosed by the involute mar- gins of the lemma; anthers ca. 1.5 mm long: ovary glabrous. Caryopses not seen. Distribution. Guatemala and Mexico. Espejo- Serna et al. (2000) cited Trisetum pinetorum for Chiapas, Mexico. Comments. The above description is based only on study of the type specimen at US and the orig- inal description. However, the Latin paragraph of the original description by Swallen (1953) men- tioned 3-flowered spikelets, and the English para- graph mentioned 2-flowered spikelets. Spikelets with 3 florets were not found in the specimen ex- amined. Though the subspiciform panicle of T. pi- netorum resembles the species of Trisetum subg. Trisetum sect. Trisetaera, this species has a lem- matal apex with four setae, and the awn is borne on the middle of the lemma. 14. Trisetum spellenbergii Soreng, Finot & P. M. Peterson, sp. nov. TYPE: Mexico, Chihua- hua: Mpio. Ocampo, Parque Nacional de Cas- cada Basaseachic, 2100 m. on nearly barren rock at overlook ca. 1 km S of Cascada, З Oct. 1986, R. W. Spellenberg, R. J. Soreng. R. Cor- ral & T. Lebgue 8654 (holotype, US!: isotypes, ESAHE!, ID!, К!, MEXU!, MO!, NMC!, RSA!, SI!, TAES!). Figure 4. Planta rhizomatosa; culmi 40—60 em alti, glabri; vagi- nae барат puce ei ше E superio- ae; laminae 20-30 ст X 2-3.5 Ф YD 7-9 mm longae, 900 ee florae; glumae subaequales; gluma inferior 5.5-6.5 X 0.3-0.5 mm, quam superior paulo bre- vior, l-nervia; ann ma superior 05-7. 5 X 0.6-0.8 mm: lemmata glabra, 5-nervia, apice Майпо; arista 8-12 mm longa, ds medio inserta; callus pilosus: palea 3.5-5 mm longa, quam lemma sua brevior; lodiculae ad apicem bi- lobulatae, ciliatae; stamina 3; ovarium glabrum. Perennial. with short rhizomes; culms 40-60 cm 24 Annals of the Missouri Botanical Garden IV Ñ ARTANGERIM 2004 lis E f. \ \ Figure 4. Trisetum spellenbergii (Spellenberg. Soreng, Corral & Lebgue 8654). —A. Habit. —B. Inflorescence. — C. Sheath, ligule, and portion of the blade. —D. Spikelet. —E. Floret. —F. First glume, dorsal view. —G. Second gul | | g glume, dorsal view. —H. Lemma, dorsal view. —I. Palea, ventral view. —J. Palea. dorsal view. К. Palea. lodicules. and pistil. —L. Lodicules (2). —M. Stamen. —N. Pistil. Volume 91, Number 1 2004 Finot et al. 25 8 Peyritschia & Sphenopholis tall, geniculate at the base, glabrous; nodes 2 to 4, glabrous. Leaf sheaths glabrous, longer than the in- ternodes, the lower sheaths loose and pilose, the upper ones open only at the apex; margins mem- branous sometimes on one side projecting upward as a sheath-auricle as long as the ligule: ligule 4.5— 6 mm long, scabrous on the dorsal surface: apex deeply lacerate, without cilia: blades 20-30 cm х 2-3.5 scabrous above. mm, flat, soft, glabrous on the lower surface, with or without a few long tri- chomes on the upper surface: uppermost culm leaf blade 10—20 cm long. 2-0 cm. lax. open, exserted; branches in 3 to 5 whorls of 2 "unicles 9-L to 5 branches per node, each branch bearing spike- lets to near the base; lower branches 3.5-5 cm long. Spikelets 7-9 mm long, (4)5- or 6-flowered: rachilla ca. 1.5 mm long, pubescent, the trichomes 1-1.5 mm long, the rachilla prolonged beyond the uppermost floret bearing a reduced floret; glumes 5.5-7.5 mm long, shorter than the spikelet, green, purple to the apex and margin; keel smooth or slightly scabrous; first glume 5.5-6.5 mm long, slightly shorter than the second, glume 6.5-7.5 mm long; lemmas 4-6.5 mm long, |-nerved; second glabrous, conspicuously 5-nerved, awned; inter- mediate nerves not extended as apical awns; mar- ginal nerves vanish below the apex: apex bilobed, the lobes each ca. 2 mm long; awn 8—12 mm long, borne on the middle of the dorsal surface, twisted at the base. geniculate, usually twice as long as the mm lemma: callus pubescent, the trichomes ca. long: paleas 3.5-5 mm long. shorter than the lem- ma. 2-nerved, the nerves scabro-ciliated; lodicules 1-1.5 mm long. the apex entire or with 2 short and unequal lobes, ciliate: anthers 1.2-2.5 mm long: ovary glabrous. Caryopses not seen. Anatomy and micromorphology. Sheath ligule apex is composed of long cells with a few very short trichomes and the epidermis with prickles princi- pally on the lower half and margins, no trichomes or stomata seen. Leaf blades in transverse section keeled. with prominent adaxial ribs, furrows be- tween ribs more than one half the thickness: scle- renchyma of first- and second-order vascular bun- dles with adaxial and abaxial girders: adaxial girders ‘T-shaped, abaxial girders [-shaped: margin- al sclerenchyma present: epidermis with costal/in- tercostal zones differentiated, the intercostal zone only with long cells more or less rectangular with straight side walls. and costal zone with cork cells and silica bodies in short rows: adaxial epidermis with short trichomes, bulliform cells inconspicuous: abaxial epidermis with hooks, stomata present only on the adaxial surface. Distribution. Trisetum spellenbergii is endemic to Chihuahua and Sonora, Mexico. Comments. Trisetum spellenbergii is morpholog- ically similar to T. viride but can be separated from the latter species by the presence of rhizomes (vs. absent in J. viride), its shorter stature (culms 40— 60 cm tall in 7: spellenbergii vs. culms 50—150 cm tall). soft and glabrous blades (vs. stiff and glaucous blades). spikelets with a greater number of florets (4- to 6-flowered in T. spellenbergii vs. 2- or 3-flow- ered), and lemmas with long bilobed apices (the lobes ca. 2 mm long in T. spellenbergii vs. less than 0.5 mm long and bidentate). Etymology. specific epithet honors Ri- chard William Spellenberg (1940—) a Mexicophile and plant systematist at New Mexico State Univer- sity. 0. Sonora: Sierra т Occiden- ward Yepachic, 1992, P. M. Раво МЕХІС tal, 9.6 mi. Е of Maycoba on Hwy. 16 t To m, 28?26' 13"N, 108?33'05"W, 14 b Peterson & C. R. Annable 12509 (US 15. Trisetum tonduzii Hitchc., N. Amer. Fl. 17(8): 558. 1939. TYPE: Costa Rica. Mount Poás, 2460 m, Nov. 1896, A. Tonduz 10749 (holotype, US-358711!). Perennial with rhizomes; culms 30-85 cm tall, erect, glabrous. Leaf sheaths glabrous, usually with trichomes at the apex, with one of the sides ex- tended up as a sheath-auricle; ligule 1-3 mm long, truncate, dentate, ciliate; blades 3-21 cm X 2-6 mm, flat or involute, stiff, glabrous, sae | loosely pubescent above. Panicles : 2-10 cm, lax, open, ovoid-pyramidal, with E us gla- brous branches, usually as long as wide, purple: rachis glabrous; branches up to 8 cm long, densely flowered above, verticillate, up to 7 per node. Spikelets 4.5-6.5 mm long, 2-flowered: pedicels elabrous, scabrous distally; rachilla 1-2 mm long. pubescent, the trichomes 3 mm long: glumes 4-6.5 mm long, equal to subequal, nearly as long as the spikelet. usually purple, acute or awn-tipped, sometimes the first glume slightly shorter and nar- rower than the second glume; first glume l- to 3- nerved; second glume 3-nerved; lemmas 3.8-6 mm anceolate, glabrous, awned, slightly scabrous, long. 5-nerved, the nerves conspicuous toward the apex. the intermediate and marginal nerves prolonged be- yond the apex as 4 setae or teeth, the setae 0.6—1 mm long; awn 7—13 mm long, borne a little above the middle, bent and twisted: callus pubescent. the trichomes 1.2-1.7 mm long; paleas 2.8—5.7 mm long. slightly shorter than the lemma, free from the lemma: lodicules truncate at the apex: anthers 1—2 Annals of the Missouri Botanical Garden mm long, purple; ovary glabrous. Caryopsis 2 mm long: endosperm soft. Chromosome number. п = 14 (Pohl. 1980). Anatomy. Leaf blades in transverse section are without well-developed keels or well-developed ribs, furrows more than one half the leaf thickness, bulliform cells in groups of 4 or 5 arranged in fan- shaped groups; trichomes absent in both epider- mes; vascular bundles with adaxial and abaxial girders, except the marginal bundles; marginal sclerenchyma present: stomata present on the ad- axial epidermis only; abaxial epidermis with inter- costal rectangular long cells, costal short cells sol- cork present intercostally. itary or cell-silica body pairs, prickles Distribution. Costa Rica and Panama, 1500— 3500 m Specimens studied. COSTA RICA. Cartago: 7 Aug. 1966, Dat iil & Pohl 842 (MO); Cantón = Orea 'amuno, P. Cordillera Central, °58'49"N, 1995, Morales & wd hin 4801 CR): Crater of Vole ‘an Irazú, 7 Aug. 1966 Pohl & Cal- ), Pittier 13082 (US); Sommet du Poas. E 841 (US): Potrero ie l alto, Volcán Poas, Nov. 1896, eie 214 (US); Sommet du Poas, 1 (P). San José: Las Nubes, 20-22 Mar le "38398 (US). PANAMA. Chiriquí: Chiriquí. 18 Mar. 1938, White 54 (MO) 16. Trisetum viride (Kunth) Kunth, Revis. Gra- min. 1: 101. 1829. Basionym: Avena viridis Kunth, Nov. Gen. Sp. 1: 147. 1816. Triseta- rium viride (Kunth) Poir, Encycl. 5: 366. 1817. Deyeuxta viridis (Kunth) E. Fourn., Bull. Soc. Bot. France 24: 181. 1877. TYPE: Mex- ico. "Crescit in alta planitie Mexicana inter Salamanca et Queretaro, 900 hexap.. Hum- boldt & Bonpland s.n. (holotype, Pl: isotype, US-865584 fragment ex P!). Trisetum bosse Swallen, BD de 4: 423. 1953. TYPE: Guatemala. Sierra de las as, E of Fine а Piamon- E P rogreso, 11 Feb. 1952. J. A. Steyerr mark US-1935005! кА US 13836 (holotype, 2208€ 306! ). Desc joue mexicana Swallen, Bol. Soc. Bot. . 1958. Trisetum mexicanum (Swallen) 5 m. 28: 233. 1979. TYPE de Bravo, 1800 m. 21 Nov (holotype, US-2119860!; iiie. MEXU not seen). Me xico 23: Perennial, caespitose: culms 50-150 em tall: nodes 4, glabrous. Leaf sheaths shorter than the internodes, glabrous or pubescent, ciliate at the apex; ligule 0.5-1(-2) mm long, membranous; apex 1 X 2-6(- usually flat, somewhat stiff, sometimes con- obtuse, minutely ciliate; blades 14—35 « 9) mm, volute, glabrous with the adaxial face glaucous, the abaxial green. Panicles 12—30(—37) X 2-8 em, lax, open or somewhat contracted, greenish to yellow- ish; lower branches 5-16 cm long, ascending, ver- ticillate; rachis glabrous. Spikelets 5-8 mm long, N - or 3-flowered; pedicels 1-3 mm long, glabrous; rachilla 1.5 mm long, pubescent, the trichomes 1.5-3 mm long especially near the apex, the lower portion subglabrous or pilose; glumes 3-7 mm long, subequal, somewhat translucent; first glume 3-6 mm long, l-nerved, slightly shorter than the spike- let; second glume 4—7 mm long, 3-nerved, equaling or slightly longer than the spikelet; lemmas 4—7 long, glabrous, awned: bidentate, 5- mm apex nerved, the intermediate and marginal nerves sometimes prolonged as 2 or 4 hyaline setae ca. 0.2 mm long; awn 5—10 mm long, borne from the mid- dle twisted: to the lower third fourth, geniculate and callus obtuse, pubescent, the trichomes 0.5-1 mm long: paleas 3-6 mm long, shorter than the lemma, hyaline, 2-nerved, the keels scabrous: anthers 1—3.3 mm long; ovary glabrous. Caryopses 2-2.5 mm long; endosperm liquid. Chromosome number. 2n = 28 (Herrera-Arrie- ta, 200 Anatomy and micromorphology. ligule apices have papillate cells and short trichomes; epidermal long cells elongated perpendicular to the vertical axis of the ligule, rectangular, with undulate side walls; no trichomes, prickles, or stomata were seen in the epidermis. Leaf blades in transverse section are without a well-developed keel but have well- developed ribs, furrows more than one half the leaf thickness, trichomes present only in the adaxial epidermis; first- and second-order vascular bundles with both adaxial and abaxial girders, the marginal bundles with sclerenchyma in small abaxial strands; abaxial epidermis with intercostal rectan- gular long cells with undulate side walls, costal short cells present as cork cell-silica body pairs, sometimes solitary; stomata and trichomes not seen, prickles present. Distribution. Trisetum viride grows in Mexico and Guatemala between 1500 and 3100 m. In Mex- ico it occurs in the following states: Chihuahua, Chiapas, Durango, Guerrero, Guanajuato, Jalisco, México, Michoacán, Oaxaca, and Querétaro (Her- nández-Torres & Koch, 1988; Espejo-Serna et al., 2000; Herrera-Arrieta, 2001) Comments. | Hernández-Torres and Engleman (1995) more recently performed an anatomical sur- vey of Trisetum in Mexico and they considered T mexicanum a distinct species. They noted that T. mexicanum could be distinguished by the size of Volume 91, Number 1 2004 Finot et al. 27 Trisetum, Peyritschia & Sphenopholis the bulliform cells, but no further morphological characteristics were discussed. We choose to place T. mexicanum as a synonym of T. viride. Specimens studied. MEXICO. Coahuila: Sierra Madre Oriental, ca. 5 km E of Saltillo (Las Palapas) up camino de Quatro, 20 Sep. 2003, Peterson et al. 17863 ( ИН. US). Durango: along Mazatlán-Durango hwy.. 3-15 kn toward El Salto from the Sinaloa iex at E | Palmito, 3 Apr. 1965, Mc p е km № carretera Durango-Mazatlán, 6 € 1978. Ed & Sánchez 78155 (US): San Ramón, 21 pend FE 1906. Palmer 128 (US): Sierra pe Occidental, 2.4 mi. N of о N of hwy. 40, 25 Oct. 1995, Pot "lerson et E 13. 2 mi. N of Tepehuana on road toward Mezquit on 2003. Peterson & E Sánchez-Alvarado 17739 * m US) Guerrero: entre Avusinapa y Petatlán, 1894, Nelson 2123 (US); Teotepec, 11 May 1939, BÉ et al. 14795 (US). Oaxaca: Sierra de San Felipe. 18 Sey 1894. Pringle 4919 (P. US). 17. Trisetum virletii E. Fourn.. Mexic. Pl. 2 108. 1886. TYPE: Mexico. San Luis Potosí: 1851. Virlet 1304 (lectotype, designated by Fi- not (2003b: 675). Р!: isotype, US-91219 frag- ment ex P-Fourn. 247!). ae PI. 2: 108. 1886. lexico. In sylva de la desierta Vieja vallis 1865. E. Bourgeau 1304 (holo- type, Ply 1 7H not seen, US-fragment ex P- Fourn-248!; US- еш nt ex MPU-Fourn-248!). Trisetum bambusiforme E. Fourn.. TYPE Perennial with rhizomes: culms 1.5-3 m tall, 5— 1(-10) mm diam. below with extravaginal branches on the upper nodes, glabrous, bambusiform. Leaf sheaths minutely scabrous, longer than the inter- nodes; ligule (2—)5—10 mm long, membranous, apex obtuse to truncate the cilia . laciniate. and ciliate, up to 2 mm long; blades 30-45 cm X 5-13 mm, flat. minutely scabrous. Panicles 20-45 х 5-12(- 20) em. lax, + yellow-green; branches usually 10-15 em а long, ver- open, pyramidal, dense, green or ticillate: rachis glabrous. Spikelets 5-7.5 mm long. (3)4- or 5-flowered: pedicels 1—4 mm d sca- brous; rachilla 0.5-0.7 mm long. densely pubes- cent, the trichomes 2—3.3 mm long, more numerous at the awned, the first slightly shorter and narrower than apex; glumes unequal, acute to shortly the second, both glumes shorter than the spikelet, scabrous along the nerves; first glume (3—)4—5.1(— 6) mm long, linear-lanceolate to lanceolate, 1- 4—6(—8) mm long, lanceolate 3-nerved; lemmas 4—6 mm long. scabrous, 5-nerved, awned, the nerves not nerved; second glume to. ovate-lanceolate, conspicuous near the apex, somewhat rounded in cross section, the margins hyaline toward the apex: apex 2- or 4-dentate (sometimes appearing almost entire), the teeth short not prolonged as setae or awns; callus obtuse with a few short trichomes, the trichomes 0.5-1 mm long: aun 1.5—4(—4.5) mm long. borne on the upper third, geniculate, slightly twisted: paleas 3.5—5 mm long, shorter than the lemma, 2-nerved, the nerves scabrous; anthers 1.8— 2.5(-3) mm long. Caryopses 1.5-2.5 mm long: en- dosperm liquid. Chromosome number. 2n = 28 (Tateoka, 1962). Anatomy and micromorphology. ligule apex has long cilia and short hair: epidermis composed of long cells with undulate side walls, with abun- dant prickles; no trichomes or stomata were seen. af blades in transverse section are without a well- developed keel or ribs, furrows less than one half the leaf thickness with trichomes present on the adaxial epidermis only; vascular bundles with ad- axial and abaxial girders; marginal sclerenchyma as a small rounded strand, stomata present in the adaxial epidermis only; bulliform epidermis pre- sent; adaxial epidermis with rectangular long cells with straight side walls, short costal cells present, stomata absent, prickles present. Distribution. Endemic to Mexico (Chiapas, Dis- trito Federal, Durango, Guanajuato, Jalisco, Méxi- co, Michoacán, Morelos, Puebla, Querétaro, San Luis Potosí, and Veracruz (Espejo-Serna et al.. 2000; Herrera-Arrieta, 2001; Finot, 2003b). Trise- tum virletii grows in moist woods between 2000 and 3500 m. У аце: Э See d. MEXICO. Distrito Fede ral: C er- ro Gordo, Cerro Cautillo Grande, Station, бит N and W of old hwy. 95, \ Iltis, Koeppen & F. Iltis 152 (US). Hidalgo: along Hwy. 105 between Pachuca and Tampico, Же inity of del Monte, 20°08'N, 98°3'W, 27 Feb. 987, non 65794 (Sl ex MO). Jalisco: Volcán Colima, 19*38'N, 103%35'W, 31 Dec. Guzmán & Nee 1065 (US); SE NS of Nevado de Coli- ma, 2 Apr. 1951, McVaugh 11721 (US): N slopes of Ne- vado de Colima, 2 Apr. 1949, m a 10161 (US). Méx- ico: Mpio. Amecameca, cañada del cerro Venacho, 12 km al E de Amecameca, 23 Dec. 1976, Koch 76337 (US); vicinity of La Cima railroad opine on top of the Serjana e Ajusco, 19°07'N, 99*12'W, 2 1963, Iltis Я Iltis 1671-a (US); Valle de Mexico, С. Tb ps 7 Jan. 1 20892 (US); pm Cerro Venacho, $ Feb. 953, Matuda et al. 28076 (US); Sierra de Las Cruces, Я Арг. 1898, 3 zh (US); Ixtaccihuatl, p 1906, Purpus 1612 (US); Temascaltepec, Mesón Viejo, 12 Jan. 1932, Hinton 2745 (US); "ин ca. Cerro Texaltepec, 3 km SSE de San Pablo Ixayoc y 13 km SE Texcoco, 9 Nov. 1975, Koch 75706 (US). Michoacán: vicinity of Morelia, 9 Feb. 1911, Arséne 5410 (US); cool slopes of mountains near Patzcuaro, 21 Dec. 1891, Pringle 3970 (US); Coal- comán, Barroloso, 3 Mar. 1941, Hinton 15747 (US): Mt s itaro, 22 July 1941, Leavenworth & Leavenworth 1122 dedere e Tres Marías, 16 Dec. 1907, Pringle 15002 1995 5 mi. S of Morelos-Federal District border, 14 Mar. 1961, pee 4155 (US) Mineral С roat & Han- Annals of the Missouri Botanical Garden Literature Cited Beal, W. J. 1896. Holt., New York. Clayton, W. D. & S. A. Renvoize. 1986. num. ae of the world. Kew Bull. Add. $ Clebsch E. C. 1960. € ;omparative Morphologic al aa Bd Variation in Arctic and Alpine. Popula- Grasses of North America, Vol. II. H. Ge nel ra кеш tions of Trisetum spicatum. Ph.D. Dissertation, Duke University, Durham, hg : 8 ina Davidse, G., M. Sous “С ink 1994. Flora Me- soamericana, Vol. 6. in smataceae a С 5 versidad Nacional Autónoma de Méxic P México, D.F Missouri Botanical Garden, St. The Natural His- tory Museum, London. Edgar, E. 1998. Trisetum Pers. (Gramineae: Aveneae) in ew Zealand. New Zealand J. 53€ Erdman, K. 8. 1965. Taxonomy of EN genus Sphenopholis (Gramineae). lowa State . Sci. 39: 289— 336. Louis; Bot. 36 564. s Monoc otiledoneas Me xica- nas. Poaceae Barnhart. Uns Sinopsis Florística. Partes IX En Finot, V. L. 2003a. Peyritschia. P. 478 in R. J. Soreng, Р. M. E erson, G. Davidse, e J. Judziewiez, F. O. Zuloa- ga, T. S. Filgueiras & O. Morrone (editors), Catalogue of New World Grasses NA eae): IV. Subfamily Pooi- deae. = U.S. Natl. Herb. 48: 1— . 206 Tse Pp. 059-076 in R. J. Soreng. P. M. Peterson, С. Davidse, E. J. Judziewicz, F. O. Zu- loaga, T. S. 5 & O. Morrone (editors), Catalogue of New World Grasses ied eae): IV. Subfamily Pooi- ea Natl. Herb. 48: 1—730. . Mexicanas Plantas ll. ae 1-10. Pari Gould, F. ге & R. Moran. 1981. The grasses of € ifornia, México. Mem. San Diego Soc. Nat. Hist. 12: 1— 140. > deae. Contr. — bs 18€ Gramineae l— Hernández-Torres, I. & E. M. Engleman. 1995. Anatomía a lámina falla del género Trisetum (Gramineae: Pooideae) en Acta Bot. Mexicana 31: 39—50. & S . D. Koch. 1987. The status of the genus Pey- de ÉXICO. ritschia (Canina ae: Pooideae). Phytologia 61: 453- 155 . 1988. Revisión taxonómica del género € Trisetum (Gramine ae: 1 ') en México. Agrociencia 71: 71-10 Herrera- Arteta. Y. 2001. Las Gramíneas de Durango. Co- misión Nacional para el Conocimiento y Uso de la Bio- diversidad (CONABIO) e Instituto Politécnico Nacional. nin —— ——— & J. Rzedowski. 2001. Gramineae. Pp. 999-1114 in G. к нын de Rzedowski & J. Rzedowski (editors), Flora Fanerogámica del Valle de México. Instituto de Ecología, A.C. y Comisión Nacional para el Conococi- aj: nto y uso s la Biodiversidad (CONABIO), Pátzcua- . Mic h loac Hit he 'ock, A. 8. 1908. Types of American grasses. Contr. ‚ Natl. Herb. . 1913. Mexican grasses in the United States Na- tional He edu, Contr. U.S. Natl. Herb. 17: 181—389. 1915. Nev Amer. J. Bot. 2: 299-310. . 1927. New species of аа from Central Amer- Рик, Biol. Soc. Wash. 88. w or note worthy grasses. ica. . 1935. Manual of the Grasses of the United States, Ed. l. ds 5. D. A. Misc. Publ. 200. 239. аа dee Scribn., Trisetum Pers., M and йч ча Fourn. Pp. 544—548, 55 A. 1-561 in Howe, H. A. Gleason & T H. Barnhart (editors), North AM n Flora 17(8): 543-638. New York Botanical = т, E — ‚С m 1951. Manual " the ( U er B Ed. 2. U.S.D.A. Misc. Pub is п, E. 1959. The 17 cum complex. Svensk Tidskr. 53(2): 203-228. Jone n B. 1980. ien Pe rasses of the 1. 200. rs. Fl. Europaea 5: 220-224. S. D. 1979. The d of three Mexican Ave- neae rand a new characters for dining vid champsia 5 Trisetum dr Taxon 28: 225- 235. Louis-Marie, P. 1928-1929, The genus Trisetum in Amer- i 30(359): 209-223. 1928. Rhodora 30(360): 237—245. 1929. qe rra R. 1983. Gramineae. 136. Th че calfe 2, C. 1960. Anatomy of the Monocotyledons. I. Gramineae. Oxford Univ. orde & Flora Novo-Galiciana 14: Press, London. J. Valdés-Reyna. 1983. Trisetum curvi- m (Poäceae; R a new species from México. Brittonia 35: 374-37 Pohl, R W. 1980. 5 y = eae (editor), Flora C и ensis. Fieldiana Bot. : e 1994. cnm 19 5 Pp. 233-235 in G. he М. urs Е A. О. Chater (editors), Flora Mesoamericana, Vol. Alis : nivers sidad Хас Еу ; Missouri Botanic 4 Gan Br Museum, 0058 n. — & 2002. Flora н qu sricana. (http:// mobot. age anda gi- V ae lv Randall, J. L. & K ilu. 1986. p m studies of North Aine ric ican Trisetum spicatum (Poaceae). Syst. Bot. 11: 567- len, St. Louis: The Natural Rúgolo de pi . E. & E. G. Nicora. 1988. Nuev taxones Lane la Argentina, y € hile пан Bol. Arge se . 25(3—4): 463—470. Se arias T. n L.] The genus eem Rhodora 8: 137-14 J. Judziewiez, 'ueiras & O. Morrone. 2003 s (Poaceae): IV. Subfam- ua R. P „ M. Peterson, G. Davidse, E. F. Za d T. S. Filg е кчө of New World grasses ily Pooide Contr. U.S. Natl. Herb. 48: Ste wudel, E. G. 1854. Synopsis ү манн! кезу earum. 2 vols. Part р 5 J. B. Metzler, Stuttgart. а J. R. 3. New grasses from Шеш, Colom- bia, Venezue “i Ecuador Bolivia and Brazil. Contr. U.S Natl. Herb. E 51-275. 195 yn grasses from Guatemala. Phytologia 1: 4 am. Tateoka, 1962. A cytological study of some Mexican grasses. E Torrey Bot. Club 89: 77-82. Tsvelev, N. 1970. De due ee 1 5 Pers. et е Pers. in URSS. Notulae Systematicae 7: 59-7: 1983. Grasses of the Soviet Union 17 05 SSSR]. Botanic heskii Institu im. V. L. Komarova and Smithso- nian Tee Amerind Publ., New Delhi. Veldkamp, J. Van der Have. 1983. The genus Trisetum in Mele sia and Taiwan. Gard. Bull. Singapore 30: 125-135. Watson, L. & M. J. Dallwitz. 1992 (onward). The Families of Flowering Plants: Descriptions, Illustrations, Identi- Volume 91, Number 1 2004 Finot et al. 29 Trisetum, Peyritschia & Sphenopholis fication, and Information Retrieval. Version: 14th De- cember 2000. анге ойнен eee APPENDIX I. Subgeneric treatment of Trisetum in Mexico and Central Americ Trisetum Species, incertae sedis: J. angustum, T. filifolium, T. pringlei Trisetum subg. Trisetum Panicles spiciform or more rarely contracted to open. Spikelets usually 2- or 3-flowered: glumes lanceolate, un- equal 1 ia or E iib : first glur . shorter or longer than the e 1(3)-nerved, usually shorter and nar- rower e the. sec uen second glume 3(5)-nerved; lemmas laterally lanceolate, 1 or pilose, slightly keeled, = 5 = = Y Y E e л a , the nerves к б beyond the awned from the upper third: awn оа a Даш or merely divaricate. some- glabrous or with short and aes trichomes at the apex. brous or with short trichomes at the apex: endosperm La. uid or more rarely solid. 2-dentate apex as 2 ог. Caryopses compressed, soft, gla- Trisetum subg. Trisetum sect. oe Species: T. curvisetum, T. irazuen Trisetum subg. Trisetum sect. сен taera Asch. & Graebn.. Syn. Mitteleur. Fl. 2: 270. 1895. TYPE Trisetum spicatum (L.) К. 9 Pl. Eur. 1: 59. 1890. Panic spiciform, narrow, densely flowered: rachis densely pilose to pubescent below the inflorescence Spikelets 2- to (2)-flow E glumes unequal, ads shorter than the us let, some- owered, rarely l- or limes ogee Species barbatipaleum, T. ligulatum, T. rosei, T. spi- catum Trisetum subg. Deschampsioidea RP phe Finot, 0 stat. nov. Basionym: Trisetum subs d sioidea Louis-Marie, Rhodora 30(5 o 928. ТҮРЕ: Trisetum palmeri Hitchc., b кз» i ed here. Panicles lax, open, rarely spiciform. Spikelets 2- to 6- flowered; glumes 2, + equal, lanceolate. keeled, muticous, glabrous, pre shorter than or equaling the spikelet: first glume 1-nerved; second glume 3-nerved: lemmas lan- ceolate, membranous, with the apex and margins hyaline, glabrous or rarely hirsute, slightly rounded in cross sec- tion, 5-nerved, the nervy pearing toward the apex or rarely н ae the 2-dentate apex in (2) or 4 setae, usually aw from the middle: awn geniculate; ovary usually оо or rarely with short trichomes at the apex soft, usually glabrous, rarely with short and shining trichomes at the apex. Species: T. di meri. 1. pinetorum. T. virletii Comments. & lectotype was chosen because Louis- Marie (1928) cited two species without designating a type. тез disap} Caryo ses compre ssed, irangense, T. martha-gonzaleziae, T. pal- spellenbergii, T. tonduzii, T. viri а & APPENDIX 2. Li: names are eoa ad in bold and synonyms are italicized. st of names and synonyms. Accepted sión Besser ex Schult. & Schult. f. = Trisetum Agrostis obtusata (Michx.) Steud. = S. obtusata Aira koelerioides de = P. тае Aira mexicana Trin. Aira obtusata Mic hx. tusata ; uxioi Avena == Kunth = S. interrupta а inter- ta е ll Kunth = T. mm Arena у J. Pres] = P. deyeuxioides Avena viridis = T. vir = Calamagrostis irazuensis Kur = Ts i iraz po Buckley — . s. inte NUM subsp. interrup Desc Dn Loslerioidts (Pevr.) Benth. = P. koelerioi- des Deschampsia mexicana Swallen — T. es ое my у d = P. pri olut mn P. 8 1 grac ilis Г B ра = Р, humilis Deyeuxia triflora Nees = eyeuxioides veuxia viridis (Kunth) E. Fourr A de Eatonia annua Suksd. = S. Eatonia densiflora E. Sn = S. obtusata Eatonia obtusata (Michx.) A S. obtusata Eatonia obtusata var. purpurascens Vasey ex Rydb. & Shear = S. ob usa Eatonia кила уаг. к ИЯ Vasey ех L. Н. Dewey = S. bt obtusa “e ola var. robusta Vasey ex Rydb. = S. ob- Eatonia 5 Scribn. & Merr. = S. obtusata atonia p (Vasey ex L. H. Dewey) Rydb. = S. ob- do ben 8 E. Fourn. = P. koelerioides glei Scribn. ex Beal = T. pringlei ob ae a Trin. ex = = S. obtusata Koeleria paniculat a Nutt. = S. obtusata Koeleria spicata Rehb. ex a & Lange = T. spicatum Koeleria subspicata (L.) Rchb. = T. spicatum coeleria truncata (Muhl.) Torr. = S. ob Peyrits oe К. Fourn. Peyritschia deyeuxioides (Kunth) Finot a FT humilis (Lou ي‎ not Peyritschia koelerioides (Peyr.) E. Peyritschia pringlei (Se 'ribn. i S. D. Koch Poa obtusata (Michx.) Link — S. obtusata Rebentischia Opiz — 7 set Reboulea gracilis Kunth — S. obtusata Reboulea obtusata (Michx.) A. Gray = S. obtusata ша obtusata var. pubescens (Scribn. & Merr.) obtusata Fe strina qoe = um мк pubescens рак = Т. spicatum olis Scrib Farw. n dou 155 annua (Suksd.) A. Heller = S. obtusata . ени » ibn.) Scribn. = S. interrupta rup Sphenopholis i interrupta (Buckley) Seribn Sphenopholis interrupta (Buckley) Saiba subsp. in- Sphenopholis interrupta subsp. californica (Vasey) Scribn. Sphenopholis obtusata (Michx.) Scribn. 30 Annals of the Missouri Botanical Garden Sphenopholis bead pp lo. purpurascens (Vasey ex Rydb. & Shear) Waterf. = S. obtusata Sphenopholis ома a lobata (Trin.) Scribn. = S Sphe nopholis visa кше, pubescens (Scribn. & Merr.) cribn. = Mus Sphe порой dian var. qu (Trin.) Seribn. ex B. tob. = S. obtusata enl obtusata var. pom (Scribn. & Merr.) cribn. ex B. L. Rob. = obtusata Re uem dba 8 Merr.) A. Heller = S. obtusata Sphenopholis robusta (Vasey ex L. H. Dewey) A. Heller = S. obtusata Trisetaria airoides (Koeler) Baumg. Trisetaria deyeuxioides (Kunth) Poir. = P. deyeuxioides Trisetaria spicata (L.) Paunero = T. spicatum Trisetarium Poir. = Trisetum Trisetarium disi (Kunth) Poir. = = Т. spicatum S. interrupta subsp. in Жш bw (Kunth) Poir. = T. viride Trisetum Pers Trisetum ай. Deschampsioidea (Louis-Marie) Finot Trisetum sect. Trisetaera Asch. & Graebn risetum sect possa Trisetum subg. Trisetum Trisetum subsect. Desc hampsioidea Louis-Marie = Tri- setum subg. ha idea Desc pr Trisetum airoides (Koeler) * Beauv. ex Roem. & Schult. _ e» аша ери (Е, Fourn.) Scribn. = P. koelerioides Trisetum altum Swallen — T. viri Trisetum angustum Swallen Trisetum bambusiforme E. Fourn. — T. virletii Trisetum barbatipaleum (Hulten ex Ve rem Finot Trisetum فو بت‎ Vasey = S. interrupta subsp. ca lifornica risetum curvisetum Morden & Valdés-Reyn: Trisetum deyeuxioides (Kunth) Kunth = P. d iode Trisetum deyeuxioides var. pubescens Scribn. ex Beal = P. deyeuxioides Trisetum durangense Finot & P. M. Peterso Trisetum elongatum (Kunth) Kunth = S. interrupta subsp. pta Triseti tum dum (E. Fourn.) Hitche. = P. deyeuxioides Trisetum filifolium Scribn. ex Be: Trisetum filifolium Seribn. ex Beal var. filifolium Trenit filifolium var. aristatum Seribn. ex Beal = T. ir — >= A. z & ~ = = i = ~ 8 > = pa = = =. D = 2 a = = [3 = = = — — . : azuense Trisetum gracile E. Four. = Cr, irazuense Trisetum hallii етп. = S. interrupta subsp. inter- rupta Trisetum humile Louis-Marie = P. humilis Trisetum interruptum Buckley = S. interrupta subsp. in- terrupta Trisetum interruptum E. Fourn. = T. s Trisetum interruptum UNR: ANKE D hallii (Se dn: ) Hitche. S. interrupta subsp. interrupta Trisetum interruptum var. californicum Doai Louis-Marie = S. interrupta subsp. californica Trisetun m irazuense (Kuntze) Hitche. orres = P. pringlei obtusa artha-go las P М. Peterson & Finot Trisetum mexicanum (Swi ag an) S. D. Koch = T. viride Trisetum nivosum E. E = T. spicatum е оге, = 1 ^ viride Ше Trisetum pringlei (Se sane ех Hel Hitche. Trisetum rosei Scribn tenerum una ibn & Мег, = T.r Trisetum rosei fo. tenerum (Sc ibn: & Mer “IT. " Louis. Marie Trisetum rosel var. nuns SC 1 0 An Hitche. = T. не Trisetum scab T. irazue Trisetum spellenl 1105 I 9 Finot & n M. Peterson Trisetum spicatum (L.) K. Richt. i spicatum subsp. lene var. Hultén = T. barbatipaleu Trisetum ангор subsp. 1 (Kunth) Hultén = Т. atum бш spicatum var. barbatipaleum Hultén = batipaleum Trisetum spicatum var. barbatipaleum Hultén ex Veldkamp I. barbatipaleum Trisetum spicatum var. nivosum (К. T. spicatum Trisetum subspicatum (L.) P. Beauv. = T. spicatum Trisetum tolucense iene) Kunth = T. spicatum Trisetum tonduzii Hit Trisetum viride (Kunth) Kunth Trisetum virletii E. Fourn. rivalve Sodiro = barbatipaleum T. bar- Fourn.) Louis-Marie = REVISION OF METASTELMA (APOCYNACEAE- ASCLEPIADOIDEAE) IN SOUTHWESTERN NORTH AMERICA AND CENTRAL AMERICA! Sigrid Liede? and Ulrich Meve? ABSTRACT The revision of Metastelma (Apocynaceae—Asclepiadoideae) in southwestern North America and Central America (excluding the Caribbe var. кш and M. schlechtendalii y v" arenicola, rank (M. arizonicum кү chiapense and M var. rank. Four infimicola as synon to M. schaffi a (oe anc new sv 'e recogn o M. schaffneri. The morphology of th an) is based on field observations and study of ognized (two of these split into two subspecific units, two into three subspecific units). Two new taxa, are described. J. californicum subsp. lanceolatum) and two to ma rank (M. harbigerum Sd var. trichophyllum). One ed: Ditassa mexicana as synonym to M ym to M. шал, М. macropodum as synonym to М. palmeri, 2 North and Central American species of | f herbarium specimens. Sixteen species are rc M. barbigerum Two former species are specific taxon (M. илеп) is raised from varietal to californio E subsp. lanceolatum, M. T. multiflorum as synonym касера as well as habitats. spec ies distribution, and phenology, are desc ribed. In addition to the taxonomic descriptions, Кол: and distribution maps jd f are presented for each specie ey words: Apocynaceae, КЕНТ Ө Central America, Vetastelma, North America. Metastelma R. Br. is an entirely American genus of Apocynaceae—Asclepiadoideae, with a distribu- tion ranging from northern Argentina as far north as Texas. Its center of diversity lies in the Carib- bean (these taxa are not considered herein). No re- liable estimates on the number of species are avail- able, because the concept of the genus has been subject to unusually large changes (see below). The number of published names stands at ca. 190—1he number of species is probably around 70, with 16 found in southwestern North America and Central America (as detailed here), ca. 30 in the Caribbean (including eastern North America), and 20-25 in South America (Liede, unpublished observations). Within the Asclepiadoideae, Metastelma belongs to the Asclepiadeae, the only tribe of Asclepiadoi- deae possessing pendent pollinia (Liede & Albers, 1994). The subtribal delimitation of the large tribe Asclepiadeae (ca. 100 genera, Liede & Albers. 1994) has been subject to much debate. In the first subtribal classification for the tribe, which was based solely on corona characters, Schumann (1895) placed Metastelma with its gynostegial. but purely staminal corona in the Asclepiadinae. He distinguished /rmischia Schltdl. because of its seemingly corolline corona and placed it in the Liede (1997), in define more naturally delimited subtribes. placed Metastelma (including Irmischia) in the Metastel- matinae. after Kunze's (1997) study of floral mor- phology had supported its exclusion from Asclepi- adinae. 1997) constituted a large subtribe with members in bath hemispheres. Recent molecular work. Tauber. 2000. 2002: Rapini et al.. in prep.) has shown that the New World mem- (1997) the. Metastelmatinae s. Glossonematinae. an attempt to etastelmatinae sensu Liede ( al.. bers of the Metastelmatinae sensu Liede form a clade of their own. in close relationship to the two other subtribes st restricted to the New World, Oxypetalinae and Gon- 2003). Ortho- forms a sub- olobinae (Liede, 1997: Карт et al. sia, often confused with Metastelma, tribe of its own together with Jobinia E. Fourn. and some former “Cynanchum” species (Liede et al.. prep.). Within the Metastelmatinae and species delimitation is still difficult, mainly st . genus and WIS f ! First M. p b thank the directors p correspondents of A. ARIZ, B. К, TEX. the loan of specimens and the MO, NY. P. , UPS, US, We г pleasant companion in the field and very helpful with the ? Department of Systematic Botany, University of Bayre uth, 95440 Bayreuth, ( ulrich.meve@uni-bayreuth.de. ilso hank M. К (Washington State University) or the « ANN. MISSOURI Bor. FR. GH, HAL, HBG, K. LL. M. ‘ir patience in waiting for their return. апу return of two types. Jim Conrad has been a artw зегтапу. sigrid.liede@uni-bayreuth.de: GARD. 91: 31-86. 2004. Annals of the Missouri Botanical Garden due to the extremely diverse, but poorly known South American species. In South America, the species attribution between the florally very similar genera Metastelma and Ditassa is purely formal. placing species with a simple staminal corona in Metastelma, those with a double staminal corona in Ditassa. It is yet unknown whether reduction of the inner corona lobe has given rise to “Metastelma” species that need to be included in Ditassa, and it is also unknown whether Ditassa-type flowers have arisen independently more than once from Metas- telma-type flowers by doubling of corona lobes. In South America, Rapini (2001) recently suggested transferring two Metastelma species (M. hemipogo- nioides Malme and M. hatschbachii Fontella & Marquete) to Hemipogon E. Fourn., a genus previ- ously defined by a characteristic erect, slender hab- it with pendulous flowers, and the absence of a co- rona (Fournier, 1885). The two species combine the typical habit of Hemipogon with the corona struc- ture of Metastelma, suggesting a close relationship between the two genera. In Central America (and the Caribbean) in con- trast, delimitation of Metastelma is by far less com- plex, even though the unfortunate inclusion of almost every small American asclepiad with twining habit and pendulous pollinia in Cynanchum by Woodson (1941) has led to considerable confusion. Prior to this (Schlechter, 1899), members of Orthosia Kunth have been confused with Metastelma even though they are clearly separable by a combination of a much more regular branching pattern, adaxially gla- brous corolla lobes, and an annular corona. Metastelma has never been revised over a larger geographical scale. This revision deals with all of mainland Central America, extending north to the southwestern United States, the northern distribu- tional limit of the genus, and south to the Panama Canal, including the former Canal Zone. It is based on field observations in the U.S.A.. Mexico, and Guatemala, and on study of more than 400 herbar- ium collections from the U.S.A.. Mexico, and Central America. Previous records of Metastelma are dis- persed in various floral treatments of different geo- graphical scope and, in part, of considerable age (see below). Nevertheless, these records give a good re- flection of species diversity throughout Central America. The highest diversity is found in the Son- oran desert, where Fishbein (1998) listed seven spe- To the north. di- (1994) listed two species for Arizona, and Shinners (1964) listed two cies for the Río Mayo area alone. versity declines rapidly. Sundell for Texas. To the south, diversity declines as well. Wiggins (1980) listed two species for Baja Califor- nia. Amazingly, two species are also listed for the small island of Socorro (Levin & Moran, 1989) (all hidden under “Cynanchum”), suggesting a consid- erable capacity for long-distance dispersal. For Mex- ico as a whole, Standley (1924) listed 14 species. Further south, species diversity remains low with five species listed for Guatemala, one each for Belize Standley & Record, 1936) and Honduras (Standley, 1938a), two for El Salvador (Standley, 1925) (all hid- den under “Cynanchum”). Two records are listed for Costa Rica (Standley, 1938b), but one of them, M. filisepalum, was only attributed to Metastelma with a pan question mark and is now excluded from the genus (see excluded species). For Nicaragua, Stevens 2001) listed three taxa, M. trichophyllum (M. schle- chtendalii var. trichophyllum) and two MER a p species. Species A is identified here as M. schle- chtendalii var. schlechtendalii, in which the attach- ment of the corona along the stipe is quite variable, and species B is identified as M. pedunculare. For Panama only one species is reported (D'Arey, 1987), again under “Cynanchum.” The genus is taxonomically difficult not only be- cause of its minute flowers, but also because 5 of the 15 species recognized here form widespread and variable complexes, the most extensive being M. barbigerum and M. schlechtendalii. This fact is reflected by the large number of subspecific units: subspecies for geographically well-separated taxa distinguishable by at least one distinct morpholog- ical character, and varieties for geographically less clear-cut taxa, distinguishable on the basis of a weaker morphological character, such as the indu- mentum. Like the other large American Metastel- matinae genera, Metastelma is а comparatively young genus still in the process of speciation and radiation (Liede et al., in prep Apart from these widespread complexes, several species are more or less narrow endemics, most of the degree of endemism in Metastelma has been over- these occurring in northern Mexico. However, estimated, because several species described narrow endemics (е... M. macropoda, M. sepicola) were found to be indistinguishable from species de- scribed from another area during the present study. One species, tical to М. underlining the close relationship of the Panaman- M. infimicola from Panama, is iden- eliasianum described from Colombia. ian flora with the Colombian one. Finally. a handful of specimens exist that are ei- ther closely related species or deviate considerably from found far off the distribution area of the most the most closely related species. These specimens are discussed in the text under the most closely related taxon. but until more material becomes available, it seems premature to describe new spe- Volume 91, Number 1 2004 Liede & Meve 33 Revision of Metastelma cies based on what is probably an aberrant speci- men. The material studied points to a considerable potential for long-distance dispersal, hybridization. and introgression, possibly resulting occasionally in atypical plants or typical plants in unexpected places. It is difficult to propose a hypothesis about the relationships of the species studied, so only a few obvious relationships will be mentioned. Metastel- ma barbigerum, M. schaffneri, and M. schlechten- dalii form a complex in which some specimens show considerable reticulation of characters. Me- tastelma minutiflorum and M. latifolium probably are vicariant species with coastal and inland pref- erences, respectively. Metastelma mexicanum shows some affinities to М. arizonicum in gynoste- gium structure. TAXONOMIC HISTORY Robert Brown (1810), is based on M. parviflorum, and ini- The genus Metastelma, established by tially included only that species. During the next 130 years numerous species were added from all over the Americas, most of them corresponding to the current concept of the genus. In the Caribbean Schlechter (1899) included Amphistelma Griseb. and Orthosia Metastelma, but in Central and South America, Orthosia remained distinct. Wood- son (1941) sank all North American Asclepiadeae into only eight genera, Asclepias L., Blepharodon Decne.. Cynanchum L., Fischeria DC., Gonolobus Michx.. Matelea Aubl., Oxypetalum R. Br., and Sarcostemma R. Br. While Woodson’s (1941) con- cept was useful for the fast naming of the many asclepiads emerging during expeditions to Central South careful readily that some of the enlarged genera are highly artificial— and America, and was therefore adopted, re-examination has shown in particular Sarcostemma (Liede, 1996a: Liede & Täuber, 2000) and Cynanchum (Liede & Täuber, 2002; Liede & Kunze, 2002). For Metastelma, Woodson (1941) stated that he did not know the genus well enough to carry out the numerous nec- essary name changes, a task readily carried out in the course of the various floras published after 1941 (see above). Stevens (1988) reverted to con- sidering Metastelma as a distinct genus, though without arguing his case in detail. Like Schlechter (1899), Stevens (1988) did not consider Orthosia Metastelma. ments published in the area after 1988 (Fishbein, 998; Levin & Moran, 1989) follow this usage. Liede (1997) argued for a distinct genus Orthosia distinct from The few floristic treat- for morphological reasons, but did not propose any formal transfers. Stevens (2001) was the first to re- establish Orthosia in a flora treatment. Molecular results based on cpDNA (Liede & Täuber, 2002) are supporting Metastelma and Orthosia as distinct genera independent of Cynanchum. MORPHOLOGY OF THE SOUTHWESTERN NORTH AMERICAN AND CENTRAL AMERICAN SPECIES OF METASTELMA HABIT AND SHOOTS All Metastelma species are slender twiners that may extend over several cubic meters, spreading over bushes and small trees. As far as known, all species possess a stout woody rootstock, from which several basally often slightly woody main shoots originate. In contrast to the often ladder-like branching pattern of Orthosia, where both leaf axils sprout a side branch, branching in Metastelma is highly irregular, and side branches sprout from one leaf axil only. Shoots may be glabrous, glabrescent. or pubescent, and the trichomes can occur over the entire shoot, along two lines, or along a single line. the position of which changes at every node. Tri- chomes on all vegetative parts are simple, multi- cellular. and smooth. Indumentum terminology fol- lows Hewson (1988) throughout. LEAVES The leaves are always decussate; the blades are usually ovate to elliptic, more rarely obovate or lin- ear. They bear one to three small glandular organs. colleters (e.g.. Thomas & Dave, 1991) directly at the base of the adaxial side of the blade at the transition point between petiole and midrib. Leaves can be glabrous or pubescent. Most frequently. the midrib on the adaxial side of the blade is rather densely covered with trichomes, the margin is cil- iate. while the adaxial leaf blade is glabrous. INFLORESCENCES A single. always extra-axillary inflorescence is borne at each node of the 5 part of the shoot. The inflorescences are 2- to 12-flowered, and in most cases sciadioidal C E We- berling, 1989) or only slightly cymose. Only Me- tastelma pedunculare has a distinctly cymose inflo- rescence, usually with a single dichasium at the base. Metastelma pedunculare is also the only spe- cies in which the peduncles regularly exceed 1 em in length. Flowers. The flowers, which are always borne on a pedicel to 5 mm long. аге 5-partite as char- acteristic for the Asclepiadoideae. They are usually Annals of the Missouri Botanical Garden nectariferous, but, as far as known, unscented, ex- cept for M. stenomeres and M. schlechtendalii var. schlechtendalii, for which a sweet to very sweet scent is reported (Schipp 674 for M. stenomeres; Davidse & Brant 32667, Saunders 740 for M. schle- chtendalit). They are generally very small (corolla length not exceeding 4.5 mm), and some (M. cu- neatum, M. latifolium, M. minutiflorum) tiny (co- rolla length not exceeding 2 mm). prising display is created by the simultaneous rather sur- opening of many flowers per inflorescence and many inflorescences. In general, the flowers are ei- ther star-shaped (e.g.. M. barbigerum), or campan- ulate (M. arizonicum), or almost tubular (M. sten- The calyx is basally fused, with the lobes usually less than 1 and pronouncedly acute in M. arizonicum and M. omeres). 1.5 mm long, but to 3 mm long mexicanum. Corolla. The corolla is white, cream, yellowish, brownish, or greenish abaxially, but always white to cream adaxially. Characteristic for Metastelma sometimes densely indumented, bearded adaxial sides of the corolla lobes. Several are the usually different types of simple trichomes can be distin- guished. Smooth, hispid, downwardly directed tri- chomes are found just above the tube in the center of the corolla lobe in most species (Fig. 1c). Long, slightly verrucose and soft trichomes may form the "beard" at the tip of the corolla lobes, e.g., M. bar- bigerum var. barbigerum and M. pringlei (Fig. 14, e). Shorter, more strongly verrucose trichomes are found in the apical and marginal parts of corolla lobes of taxa without a distinct "beard." e.g., M. barbigerum var. liesnerianum (Fig. Le). Corona. The corona is of gynostegial origin, but can occasionally be fused basally to the corolla (M. parviflorum, M. mexicanum). In contrast to Ortho- sta, in which the corona is fused for at least one third of its length, it consists of free staminal parts only (Kunze, 1997). and variation between species comes about by var- The corona is always white, iable length in relation to gynostegium length, shape, and fleshiness, e.g.. in M. barbigerum, the corona is about as long as the gynostegium (Fig. la), while it is considerably longer with its parts connivent over it in M. pringlei (Fig. 1b). Point and angle of attachment of the lobes are further distin- guishing characters: in both M. barbigerum var. barbigerum and M. pringlei the corona lobes form a distinct hook at the point of attachment (Fig. la, b). while this hook is much less pronounced, e.g.. in the other two varieties of M. barbigerum as well as in M. schlechtendalii. Gynostegium. The gynostegium can be either sessile, or elevated by a stalk (Fig. la, b). In gen- eral asclepiad terminology, a stipe. a feature occa- sionally found in Cynanchum, is a stalk elevating the gynostegium, but not the corona, and a column, typical for the Asclepiadinae, is a stalk elevating both the gynostegium and the corona. In Metastel- ma, the corona can be attached to the base of the stalk (e.g.. M. parviflorum), along the stalk (e.g., M. eliasianum, М. barbigerum, М. arizonicum), or di- rectly underneath the gynostegium (e.g., М. pedun- culare, М. stenomeres). While the terms “stipe” and “column” have been used in the descriptions ac- cording to their definitions, all intermediate states exist, so that the distinction is purely formal. The outline of the gynostegium is a useful character in species identification, especially for the large and variable M. barbigerum and M. schlechtendalii complexes. Metastelma barbigerum always has a conical gynostegium with inclined anthers (Fig. 1а), while M. schlechtendalii always displays a more or less cubical gynostegium with erect anthers. In the Yucatán, this character short-stiped forms of M. schlechtendalii from М. distinguishes even the barbigerum var. veracruzense and variety liesnerian- um, in which the hook-shaped corona attachment and the corolla “beard” easily identifying the typ- ical variety are often less well developed. The guide rail, formed by the anther wings of two adjacent anthers is as long as the anther proper, except for M. schaffneri, where it is slightly shorter. It consists of an outer and an inner ridge, with the inner ridge shorter than the outer one and strongly curved (Fig. 1f). Pollinaria are minute and therefore difficult to use for identification, even though the shape of cor- puscula, caudicles, and pollinia are species-specif- ic. The lower part of the style-head possesses a Figure 1. ew; two petals removed. —b. neriani um, < up} d. Metastelma pringlei, simple, slightly verrucose trichomes on upper regions of corolla lobe surface SEM studies of flowers and flower details. — Metastelma pringlei. 5 with corona. yer regions of corolla lobe surface with smooth trichomes (ce in r) and verrucose trichomes (margins), — — . Metastelma num var. nic 0 in lateral —c. Metastelma barbigerum хат. lies- . Metastelma barbigerum var. barbigerum, detail 10 verrucose hair from median region of adaxial corolla lobe siriane 'e. _( Metastelma 'arbige rum var. barbigerum, detail of gynostegium s "а and опе pollinium with е one anther removed. ede & Conrad 3242. showing one pu with dis tal and proximal ridge (arrow) of anther : Liede & Conrad 2610; b. d: Liede & Meve 2501; Volume 91, Number 1 Liede & Meve Revision of Metastelma Yor \ d AUF UM $ NN. p م‎ * „N ae І у ¡A Annals of the Missouri Botanical Garden knob above the corpusculum and the upper part of the style-head is flat to conical in most cases. Only М. mexicanum can be easily recognized by its ob- infundibuliform style-head. icles and seeds. One follicle per flower is generally developed. A compitum (Kunze, 1991) is thus absent. Follicles are slenderly obclavate to fu- siform in shape, with a thin pericarp and without wings or other ornaments. Each follicle contains 10 to 20 seeds in an imbricate arrangement on the placenta. Seeds are ovate, oblong, or pyriform, winged or wingless, light to dark brown, glabrous or papillate, smooth, with longitudinal ridges or tu- berculate, but no species with seed trichomes has been found. CHROMOSOME NUMBERS Of the species in western North and Central America, only Metastelma barbigerum var. barbi- gerum has been counted with 2n 2 (Albers et al., 1990, as Matelea reticulata (Gray) Woodson), the chromosome number found in more than 90% of all Asclepiadoideae (Albers & Meve, 2001). HABITATS, DISTRIBUTION, PHENOLOGY, AND USES Most Central American Metastelma species dis- play a rather wide ecological amplitude and occur on various soils and in numerous vegetation types. Metastelma arizonicum subsp. arizonicum is a low- land desert scrub species, and M. californicum subsp. californicum occupies even drier habitats. Both taxa have a South or Central Mexican sister taxon living at higher altitudes and in forest habitats, open pine forests for M. arizonicum subsp. chiapense and tropical deciduous forests for M. californicum subsp. lanceolatum. Of the remaining Sonoran spe- cies, M. cuneatum and M. minutiflorum are restrict- ed to the lowlands (to 200 m), where they inhabit thornscrub communities. Metastelma latifolium and M. mexicanum prefer altitudes above 200 m (to 1500 m) and live in open forest communities. In Texas, Metastelma barbigerum var. barbigerum is commonly found in lowland areas (below 500 m), while M. palmeri prefers higher altitudes (above 500 m). In Mexico, though, M. barbigerum var. bar- bigerum is also found at higher altitudes to 2300 m, while the other two varieties are strictly lowland plants of sand dunes, rivers, and disturbed habitats. A similar difference in altitudinal preference char- acterizes the three subspecies of M. schlechtendalii. The typical variety is salt tolerant and occurs main- ly in open or disturbed locations close to the sea, but is occasionally found as high as 2000 m in Guatemala. Morphologically extremely similar is the strictly coastal disjunct variety arenicola. Va- riety trichophyllum, easily recognized by its densely hispid leaves and more floriferous inflorescences, in contrast, prefers altitudes over 750 m and open forests. Other lowland species are M. stenomeres, a distinctive Belize endemic living in open pine for- ests, and М. eliasianum, extending from Panama to the west coast of South America (Colombia). The remaining four species, M. longicoronatum, M. pe- dunculare, M. pringlei, and M. schaffneri, occur regularly at altitudes between 1000 and 2500 m, although some specimens have been found as low as 500 m or as high as 2800 m. No Metastelma, however, was found above 3000 m. The latter four species, except for M. pedunculare, which has a weedy tendency, live in open forest habitats. Very little is known about the reproductive and pollination biology in any species of the genus. Liede (unpublished) observed a small bee visiting Metastelma californicum subsp. ca 5 and Fishbein (Fishbein & McMahon 2769 observed bees, wasps, and flies TEN Sp., Halicti- dae, Sphecidae, and Diptera) on the same species. The host-plant ages by Markku Savela, (http:// et. 0 MEL) 577 1.251. ww.nic.fun m. к lists Matasielma as probable host for "A butterfly Anetia thirza insignis (Salvin, 1869) (Nym- phalidae) in Panama and Costa Rica. Fruits have been observed in all but three species (M. latifol- ium, M. longicoronatum, M. stenomeres), but not to the extent suggesting that Metastelma belongs to the extremely rare asclepiads capable of self-fertil- ization, as e.g., Tylophora hirsuta Wight (Chatur- vedi, 1988) and Vincetoxicum nigrum (L.) Moench (Lumer & Yost, 1995). The phenology data pre- sented in the species descriptions are based on col- lections of these species over their entire range (see Appendices 1, 2; an index to scientific names is provided in Appendix 3). While most species show a distinct peak of flowering at a single locality, this peak may shift with changing geographical param- eters. Also, most species continue to produce a few flowers after their peak flowering time is long past. Metastelma is not generally noted among the rec- ognized useful plants. Morton (1981: 692) noted, "Indians of Yucatán boil the root and repeatedly rinse the mouth with the decoction to heal canker sores. They warn that the decoction must not be swallowed." TAXONOMY Metastelma R. Br, Asclepiadeae: 41. 1810. TYPE: Metastelma parviflorum (Swartz) R. Br. [= Cynanchum parviflorum Swartz]. Volume 91, Number 1 2004 Liede & Meve 37 Revision of Metastelma Imp. Naturalistes Moscou Acrocoryne caribaea Turcz. basset Bartlett, Arts 44: 631. 909. TYPE: Basis mexicanum (Brandegee) En [7 Melinia n mexicana Brandegee |]. lectotype, designated here. Devas Schltr., in Urban, Symb. Апи. ^E: Dec 5 broadw ayi Schltr. qu Turez.. Bull. Soc. 5(2): 316. 1852. TYPE: A Acad. 1: 264. 1899. * — FI. mi 149. 1913. Type: Epicion ba- hamense Vie Small (Metastelma | bahamense Gr Irmisc hia “Sch Idl., Linnaea 19: 738. 1847. TYPE: /rmis- chia floribunda Schldl. Meresaldia Bullock, Kew Bull. 19: 203. 1965. Esmeraldia . Fo Qc 85 =] o ~] ج‎ » 3 R=. Sy? A, = — - E 5 57 1862 (Orchidaceae). Fourn.) Bullock = Esmeraldia stricta E. Stelmation E. Fourn., in Martius, Fl. Bras. 604): 226. 1885. TYPE: Stelmation myrtifolium (Decne.) E. Fourn. [= Metastelma myrtifolium Decne. ]. to 400 em high, subterranean Plants small, slender twiners. richly irregularly branched; organs constituting a woody rootstock: shoots herbaceous, basally often slightly woody or corky, pubescent, glabrescent or glabrous; latex white to ivory; pseu- dostipules (extremely reduced short shoots with a pair of smaller and sometimes differently shaped leaves) absent. Leaves opposite, petiolate, with 0 to 3 colleters at blade base: blades herbaceous. Inflo- rescences usually one per node, extra-axillary, 2- to 12-flowered, bostrychoid or sciadioidal, peduncu- late to sessile; floral bracts present, triangular or ovate. Floral buds elongated-conical, conical, ovoid or cylindrical; calyx basally fused, campanulate, lobes ovate, acute to acuminate; corolla rotate to campanulate to almost tubular, 1.5-4.5 mm long, abaxially white, cream, yellow, green, or brown, glabrous, adaxially white, densely covered with verrucose and smooth trichomes, arranged in spe- cies-specific patterns; gynostegial corona of free staminal lobes, white, glabrous; lobes simple. Gy- nostegium sessile or atop a stipe or a column; an- ther wings consisting of distal and proximal ridge. proximal ridge short, curved; connective appendages present, ovate or triangular; pollinarium with two pollinia per pollinarium, (sub)apically attached to the caudicles, pendulous, not possessing a germi- nation pore; upper part of the style-head depressed- conical to conical, rarely flat or elongated-conical. Follicles one, occasionally two per flower, 25—80 mm long, obclavate or fusiform, terete or obtusely deltate in cross section, apically beaked, wingless, slightly longitudinally grooved, usually glabrous, with thin pericarp; seeds 4-7 mm long, brown, winged or wingless, glabrous or papillose, smooth or with longitudinal ridges or tuberculate, without trichomes, comose. Chromosome number. 2n = 22 (Metastelma barbigerum var. barbigerum, Albers et al. (1990). as Matelea reticulata (Gray) Woodson). Basistelma was created by Bartlett (1909) to ac- commodate two species, B. angustifolium (Torr.) Bartlett and В. mexicanum (Brandegee) Bartlett, that are at first glance ill fitted in Metastelma be- cause of their long, conspicuous style-head. Fish- bein and Levin (1997) correctly included Basistel- ma in Metastelma, and recognized the two former Basistelma species as identical. As the basionym of B. angustifolium, Metastelma angustifolium Torr., is a homonym of Metastelma angustifolium Turez. (a taxon belonging to the genus Orthosia) and thus not a name available in Metastelma, Fish- bein and Levin (1997) assigned the name Metas- telma mexicanum (Brandegee) Fishbein & Levin to this species. Under these circumstances, it is log- ical to select Basistelma mexicanum as the lecto- type of Basistelma, an action not carried out by Fishbein and Levin (1997). KEY TO THE SPECIES OF METASTELMA IN THE SOUTHWESTERN U.S.A. AND CENTRAL AMERICA* la. Gynostegium sessile or almos 2 so. Flowers tiny; corolla lobes not longer than ca. 2 mm (northern Pacific slope of Mexico only). 3a. Corolla lobes puberulent with very жой trichomes (< 0.1 mm long); corona distinctly 15 555 nar, at les cu ast basally as broad as the ant neatum 3b. C х Па lobes densely covered with p trichomes (> 0.1 mm long); corona lobes + ad distingüy smaller than the бү 5 o. e =< Ф | — 3 O @ -$ e © < se ho 5b. Base af stained corona lobes at the base broader than the ps plants of c 3b. M Mexico Ib. Gynostegium with a distinct stipe. a p apiculate, shoots with one line of trichomes, — with a pedun 2.4 . Flowers larger; ELA р at least 2.5 mm SE 5a. Base of staminal corona lobes жй а than the anthers; plants of northern Мех За. М. enderly lanceolate, shoots with two lines of trichomes, inflorescences d co 9. M. i tui 6. M. LI. 3. M. californicum 1 . californicum p^ ай eto eb aii entral and sout . californicum subsp. а р Stipe as long as ће gynostegium or shorter than it. Annals of the Missouri Botanical Garden Та. Corona distinctly shorter than the gynostegium; anther appendages about twice as long as the anthers, erect; style-head with a pronounced appendage; corolla lobes apically with very short (< 0.05 mm long) verrucose trichomes (giving a papillose appearance) and a tuft of smooth, stiff trichomes at the base of each corolla lob. 8. M. mexicanum Tb. Corona as long as or longer than the gynostegium; anther appendages not longer than the anthers, inflexed; style-head without a pronounced 5 corolla lobes apically with distinct tri- chomes (at least 0.1 mm long) and smooth trichomes, if present, not 3 to a small tuft. 8a. Corona ca. 1.5 mm long, about twice as long as 7 gynostegium, including stipe, concealing the gynostegium 00 00 12. M. мй 8b. Corona shorter than 1.5 mm, 1.5 times as long as the gy noste; gium (incl. stipe) or less, not concealing the conical gynostegium. 9a. din distinctly lanceolate, suddenly broadening at the base, ce an almos istate shape o . M. a 9b. Corona never sudde nly broade ning at the base to produce e a hastate P 1ape. Ja. Corona attached along the stipe, lobes subulate and bases hardly broader than apex or slenderly ovate to lanceolate, with or г without a long extended tip, often attached to the stipe with a pronounced ho la. Corona attached to the stipe with a oe ‘ed hook, lobes longer than 1 mm and style-head longer than 0.5 mm. 12а. Stems and pedicels with a eh line of rec trichomes; calyx with a few sparse trichomes; leaves less than 3 mm wide: corolla lobes oblong or ovate, but not apically sudde nly narrowing; trichomes of the EA side of the corolla lobes apically not —— longer than laterally 13 2 V. arizonicum Stems basally often corky. € veins with tric homes; calyx lobes ery slenderly ovate, almost as lon g as the corolla lobes, but at ln reac hing half of corolla 7 ah corolla lobes adaxially lat- erally and apically with long (0.2-0.3 mm), distantly verrucose tric homes; plants of the ала U.S.A. and northern Mex- x arizonicum subsp. arizonicum 13b. Stems basally never corky, lod ins glabrous, calyx lobe ovate, not very slender and not reac cra half of corolla le 8 corolla lobes adaxially apically and laterally with short (ca. 0.1 mm), strongly verrucose trichomes; Mee of southern Mexico, Guatemala, Honduras lb. M. arizonicum subsp. chiapense 12b. Stems, нет els, and calyx glabrous, corolla lobes ovate to lanceolate, en apically rather abruy Aly narrowing; trichomes of wed adaxial side of the corolla lobes apic ally distinctly iro than laterally ко М. 5 var. een 11b. orona + - flatly attac died inda 'rneath the gynostegium, either lobes shorter ian 1 mm or style- head shorter than 0.5 mm, or both. > Ma. Pedicels glabrous; stems glabrous or with, a single line of trichomes, calyx with a few isolated trichomes on b tube — >: ›. М. barbigerum var. liesnerianum 14b. Pedicels dipai ly covered with tric hones stems always with a occasionally very broad line of trichomes; calyx with trichomes с - 2с. barbigerum var. veracruzense 10b. Corona attac hed dim ctly ат the empiece: Ibis ovate to triangular, ays without a long extended tip, never attached to the stipe with a pronounced hook. 15a. Leaves, at least those of flowering branches, mostly linear or lanceolate, rarely ovate; corona lobes not distinctly flat, oblong, considerably exceeding the fla tly cylindrical gynostegium .. М. Sin dd eaves normally ovate; corona lobes dis stinctly flat and muc h broade ra the base than at the apex, barely exceeding the slightly conical gy ай ка 15b. gium. l6a. Trichomes of the adaxial side of the corolla к! and laterally of equal length and fairly short (0.1—0.15 mm); leaves na rrowly ovate, on жакын ranches not exceeding 15 mm bu corolla usually less than 2.5 mm (rarely to 3 mm) long 10. M. palmeri 16b. Tric homes of the adaxial side of the corolla apically slightly май E terally (to 0.2 mm); leaves ovate, on cli: — s to 25 nm теа corolla to 4 mm long Ob. Stipe distinctly longer than the gynostegium. 7a. Corolla fused at least for a quarter of total length. 8a. Corona attached to the stipe immediately below gynostegium. Corona almost twice as long as the gynostegium without stipe; style-head conical; — BED = e turneri — 8 Volume 91, Number 1 2004 Liede & Meve Revision of Metastelma dioidal; floral bracts elabrous eaves slenderly elliptic or геи peduncle 3— 5 mm long: inflorescences scia- 15. M. stenomeres 5. 19b. Corona just exceeding the gynostegium: style- e 1d flat; R es ovate; к ‘le at least ) mm s inflorescences bostrychoid: floral bracts ciliate Corona not attached immediately below gynostegium. 20a. Ce to stipe: corolla (ооо for less than half of its length, lobes 18b. 4h bi ittachec M. pedunculare dense, p Be woolly trichomes: stipe stout, not more than twice as long r as the ynostegium; plants of C entral and South. America / 20b. c orona attached to the corolla: pones fused for half of its length or more. labe 8 L short pube scence : stipe stegium: plants of South 5 Corolla Е» ab basally ( 17b. (corona attached кер stipe y E» M ۹ OEE М. “Тее чыш ith ery slender, at least three times as long as the gy- DEI PL. . parv iflorun т К. М. schlechte мий 21а. Adaxial leaf surface with evenly spread trichomes; inflorese ences s usually sith more than "pina pedicels with evenly distributed trichon 21b. Adaxial le id ud e (except for midrib) 1. sc Wy htendalti var. . trichophyllum hon tric isha = ‘ences usually with less than 6 flowers; pedicels with a single line of trichomes or glabrous 22a. Inflorescences distinctly pe fie ulate, peduncles normally more than 4 mm long: ы acts and sepals ovate: corona exceeding the gynostegium by less than gynosteg length: style-head fat to slightly поа Да. М. schlechtendalii var. schlec -— 22b. Inflorescences subsessile. pedune Jes less than 4 mm long; floral bracts and sepals distinctly lanceolate: corona exc 'eeding the gynostegium by as much as or more than gynostegium length: stvle-head distinc tly реса 1. * Metastelma E illorum is not treated here in detail. but is included in the key because it is widespread in the Caribbean and Soal America, and th been applied to material of M. schlechtendalii. e name has often l. Metastelma arizonicum A. Gray, Proc. Amer. Acad. Arts 19: 85. 1883. Cynanchum arizon- icum (A. Gray) Shinners, Sida 1: 365. 1904. TYPE: U.S.A. Arizona: Pima Co., hills near Tucson, 22 May 1883, C. G. iur 20 (ho- lotype. GH not seen). Figures 2a—h, la. Metastelma arizonicum A. Gray subsp. ar- izonicum Metastelma w atsonianum Standl.. Contr. U.S. Natl. Herb. 23: 1175. 1924. Basionym: Metastelma albiflorum 5. Wat аг, Amer. Acad. Arts 24: 60. 1889, Meroe аи Griseb., Fl. Brit. W. Ind.: TYPE: Mexico diim baec E. Pa He "e diste GH not seen; isotypes. NY—21). Sonore Plants ascending, 80-250 cm high: shoots pe- rennial. basally corky, isolatedly to sparsely cov- ered along a single line with recurved trichomes, ca. 250-300 um long; internodes 1—4 cm long, ca. | mm diam. Leaves with petioles 1-2 mm long. with or 2 colleters at blade base; blades 10-20 X 1.5- 3 mm, elliptic to ovate, basally rounded to cuneate, apically acute to acuminate, marginally straight or revolute (when dry). adaxially and abaxially gla- brous, but sparsely covered on the midvein and 200-250 um long. Inflorescences always one per node, l- to 4- margins with recurved trichomes, flowered, 1 to 4 flowers open synchronously, scia- dioidal, subsessile: floral bracts 1-1.5 X 0.20.5 ). M. SOS d var. eo mm at the base, deltoid, slightly papillose and with trichomes on the surface; pedicels 1-2 mm long. sparsely covered over the entire surface with re- curved trichomes, 200-250 um long. Floral buds with im- 2.5-3 mm long, 1-1.5 mm diam.. ovoid. bricate aestivation; calyx basally fused. campanu- late, abaxially vaguely glandular. with a few sparse 0.3—0.5 mm, slenderly ovate, apically acute; corolla campanu- late, 2.5—4 mm long. adaxially with distantly verrucose trichomes, 200— trichomes: calyx lobes 2-2.5 very white, abaxially glabrous, 300 pum long, on the apical and on the lateral parts f the lobes and with downward-directed, hispid trichomes, 300-500 um long, on central part of the lobes at the mouth of the s lobes fused for ca. %4 of total corolla length, 1 mm wide. patent and ecurved, apically acute: hubs corona 1.5-2 mm long. equaling to longer than the gynostegium: lobes inflexed or apically reflexed. attached at the base of the stipe with a distinct hook-shaped base. laminar to solid, subulate. Gynostegium ca. 1.5 mm high. ca. 0.7 mm diam., on a stipe 0.7 mm long: anthers about as high as broad, deltoid, abaxially convex; anther wings ca. 500 um long. extending along the entire length of the anther, anther wings of adjacent anthers parallel to each other, in the same plane as the anther to slightly centrifugal: 1. 250 um wide, triangular, equaling the stamen in width, ~ nA connective appendages ca. 400 jum long. erect; pollinarium with corpusculum 150 jum long. more than twice as long as broad, elliptic: caudicles 80 um long, cylindrical, s-shaped. convex-concave: pollinia subapically attached to the caudicles, 200— 40 Annals of the Missouri Botanical Garden \ Ан S W | 5 DNE vy 0.5 mm 0.5 mm Figure 2. Metastelma arizonic um. m p Metastelma arizonic um subsp. arizonicum. aH Metastelma arizonicum LO „саре nse. Ha LC а, a’. . —b, b’. Flower. е, e”. Corolla lobe, ac daxia view. —d'. Corona and gynostegium. —e, . Pollinarium, —f, Po Style- he de —pg. Follicle. li. Seed. a : Steinmann & Varela 972. g. h: Neese & тыя 10076. a’: Ghiesbreght 664. b'—(': Lie da & Conrad 2559. Drawn b J. Conrad. Volume 91, Number 1 Liede & Meve 41 2004 Revision of Metastelma | ac = — ad i EN | | | 2; {4 Р fo _ oe = — — 35° — b J Е | } “умыл | © Y [tC e | | E f | ‚ 4 ® % Oo B P1 1 т m NENNEN X y | | ` > | | ) — | — N Q | | | t Cà “ч JA dae * 4 s } بت‎ = i = | І 2 : = TP 30° y э „л, A taal NI as EN ў | Mi = lS НЕ ЧЕ —R ' b^ | К L A ө м. рема 55р. arizonicum b M x yi TT G ч. =; | ca | A M.arizonicum ssp. chiapense 1” х PO, | eta dn | Y © М. mexicanum E A E AE 44 3) 4 = | — | — — Ne 7 = А A | КДУ А-А -120° -115° -110° -105° -100° -95° -90° Figure 3. Distribution of Metastelma arizonicum and М. mexicanum. 220 um long. 80 um wide. ovate in cross section, clavate: style-head white, 0.4 mm diam., 0.5 mm high. upper part 0.2 mm high, shorter than to equaling the lower part, depressed-conical to con- ical. Follicles always one per flower, pendulous, 45-65 mm long. 3—4 mm diam., fusiform, terete, apically strongly beaked, medium to dark brown, glabrous: seeds ca. 10 per follicle, 5-6 mm long, 2-3 mm wide, pyriform, medium brown, seta and aseta side tuberculate, marginally with 0.15 mm wide wing with entire margin: coma 20-30 mm long. Phenology. Collected in flower in April, June, and from August to January; in fruit from March to May and September to January. Distribution. | Mexico: Sonora; U.S.A.: Arizona; mostly on rocky slopes in desert scrub communi- 10-1000 m. Literature and illustrations. pl. 18) ties: Wiggins (1940: 92, pia ig pes ns examined. MEXICO. Pape 8.3 rd. of Arizpe on MEX Hwy. 2. J Sep. 19 (fl fr), — 96-27 (ARIZ); Arroyo К | Mentidero, iE : km S of Alamos, 25 Nov. 1993 (fl, fr), Van Devender et al. 93-1310 (ARIZ); near purs 3 on NE side of Rio Yaqui hit on S ME 1 . 15 Se )98 (f). Van De- vender & Reina 98-1152 (ARIZ), y to hus San C “ү 12 Jan. 1965 (fl), Thomas & Felger 11942 (ARIZ): ca. | mi. N of Bahia San Carlos on rd. to Algodones. 24 Nov. 1963 (fl), Felger 9579 (ARIZ): Bahía Credo E side of Morro Colorado, 28 Dec. 1966 (fl, fr). Felger 15588 (ARIZ); Bahia de s Pedro, Sep. 1954 (fl, fr), Turner ү al. 79-261 0 . 5 km E of Bahia San Pedro, 17 er ud prn 84-461 (ARIZ, ie 4 (fl, fr). Van Dev ender & Van Devender et al. 84- % (ARIZ); Guaymas 38 ‚ 1965 (fl), y xem & Felger 11785 (ARIZ): achiste about Bahia Topolo- 1 Sep. 1954 (fl. fr). Ld 1 (ARIZ, LL); Cerro del Viejo SW of Caborca, 9-11 1954 (fl. fr). Gentry 14466 (ARIZ); 23 mi. 5 of hy 26 Sep. 1934 (fl). Wiggins 7491 (ARIZ): El Baviso, ca. 2.5 km NE of Bahia San Carlos, 18 Nov. 1984 (fl), Felger & Zamulio 84-490 (ARIZ); Enseñada Chica ca. 23 km N. 39 km W of Guay- mas, | Oct. 1979 (fl, fr), Burgess 5695 (ARIZ); Ensenada runde [= Bahia San Pedro]. 11 Nov. 1964 (fl). Russell et ы oleae m уо, 23 Oct. 1934 (fl). suajaray, Rio Ma Gentry 1098 (ARIZ); Guaymas, Palmer 626 (NY); 2 km E of MEX 15 on e to Microondas Las Av vispas. 21 Sep. 1997 (fl). Reina et al. 96-286 (ARIZ); 1 km E of Los Camotes, along Los Tanques-Las Chinacas rd.. 18 Aug. San Bernardo, Rio 1994. Fishbein et al. 1878 (ARIZ): 42 Annals of the Missouri Botanical Garden Mayo, 12 Feb, 1935 (fl), Gentry 1298 (ARIZ); Sierra Cip- riano, 28 Apr. 1991 (fl, fr), Peer dp ш iip arenas 91-33 (ARIZ); Sierra Libre, ca. 2 km E of the tion with MEX 15, 2 Sep. 1996 (fl), Ботла & е ‚ 972 sup NV): Sierra Seri, Hast : епа, 17 Apr. 1‹ 74 (fl. Вени et al. 74-15 с S.A. Arizona: | SU eiie Mis., 8 Oct. МА Gould et al. 2804 (NY - Babonuiven Mts., Fi yon, 7 Sep. 1931 (0). Gilman 140 Ща dicis Mts., 19 Oct. 1945 (fl), Goodding 208-45 (NY); 3.5 mi. below Mol- ino Basin poa Santa Catalina Mts., 18 Apr. (fl, fr), Keil 432 ): Santa Catalina Mts., 21 Dec. (fl, fr), Tourney s.n. 155 Sierra Tucson, 26 May 1884 (fr), Pringle s.n. (NY, topotype): Ventana Park, E of San Mi- guel, 13 Oct. 1945 (fl. fr). Condding 116- po ак Pinal Co., Superstition Mts., px ih 206, . 20 mi. E of Phoenix, 14 Apr. 1981 (A, i Yer 4 кона 10076 (NY). County not EE ge Apr. 1884 (fl, fr), Le Roy s.n. (NY); Sonora, 15. Aug. 1884 (fl), epa s.n. (NY). guesl e 2 Ib. Metastelma arizonicum A. Gray subsp. chiapense (A. Gray) Liede & Meve, comb. et stat. nov. Basionym: Metastelma chiapense A. Acad. Arts 21: 397. 1886. Cynanchum chiapense (A. Gray) Standl & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 226. 1947. TYPE: Mexico. Chiapas: 1864— 1870, A. В. Ghiesbreght 664 (holotype, Е (neg. 05966 1)!; isotypes, GH not seen, MO not seen, NY!). Figures 2a’-f", 3. Gray, Proc. Amer. „ 5 S. E Blake, Contr. U.S. Natl. Herb. 19. 2. Cynanchum соШпит (S. F. Blake — . ^ я yerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 226. 19 d TYPE: Guatemala. Pu trail fom Los Amates to Izabal, 31 May 1919 Blake 7793 (holotype, Us. 989029!; isotypes, С н по! seen, US not seer Plants ascending, ca. 50 em high, richly basi- tonically to irregularly branched. Subterranean or- gans constituting a woody rootstock; shoots peren- nial, sparsely covered along a single line with recurved trichomes, 200-250 um long; internodes 1.5-5.5 em long, 0.5-1 mm diam. Leaves with pet- ioles 1-2 mm long, with | or 2 colleters at blade base; blades 15-35 X 1.5-3 mm, linear, elliptic, or ovate, basally rounded to cuneate, apically acute to acuminate, adaxially and abaxially glabrous, but marginally sparsely ciliate. Inflorescences one, oc- casionally two per node, 3- to 7-flowered, З to 5 flowers open synchronously, sciadioidal, sessile to subsessile; floral bracts 0.6-1 X 0.3-0.5 mm at the base, ovate; pedicels 1-3 mm long, sparsely covered over the entire surface with recurved trichomes, 100-150 jum long. Floral buds 1.8-2.2 mm long, 1-1.5 mm diam., ovoid; calyx basally fused, rotate to campanulate, abaxially vaguely glandular and with a few sparse trichomes; calyx lobes 1—1.5 * ca. 0.5 mm, ovate, apically acute; corolla campan- ulate, 2-2.5 mm long, abaxially and adaxially white, abaxially glabrous, adaxially with verrucose trichomes, ca. 100 um long, on the apical and on the lateral parts of the lobes and with smooth, downward-directed trichomes, 300-400 um long, on the central parts of the lobes; lobes fused for ca. % of total corolla length, 0.8-1.2 mm wide, erect, apically recurved, ovate, apically acute; gynostegial corona 1.2-1.5 mm long, equaling the gynostegium; lobes slightly inflexed, attached at the base of the stipe with a distinct hook-shaped base, laminar to filiform, oblong. with entire margins. Gynostegium са. 0.8 mm high, ca. | mm diam., on a stipe 0.6 mm long: anthers about as high as broad, deltoid, abaxially convex; anther wings ca. 300 jum long, anther wings of adjacent anthers parallel to each other, centrifugal; connective appendages ca. 400 um long. ca. 300 um wide, ovate, equaling the sta- men in width, slightly inflexed; pollinarium with corpusculum 150 jum long, more than twice as long as broad, elliptic: caudicles 70 um long, cylindri- cal, slightly s-shaped, convex-concave; pollinia api- cally to subapically attached to the caudicles, 200— 220 um long, 80 um wide, ovate in cross section, clavate; style-head white, 0.5 mm diam., 0.5 mm high, upper part 0.25 mm high, depressed-conical to conical. Follicles and seeds unknown. Phenology. Collected in flower in April, May. July, and from September to November. Santa Ana; Guate- mala: Péten; Honduras: Morazán; Mexico: Chiapas: Distribution. El Salvador: rocky slopes, very open pine forests; 650-1700 m. Metastelma arizonicum, M. barbigerum, and M. pringlei form a group of closely related species. They share bearded corolla lobes, the conical shape of the gynostegium, and the protruding anther wings. Yet, they are relatively easily separated by the length of the corona lobes (ca. 1.5 mm long, about twice as long as the gynostegium in M. prin- glei, shorter than 1.5 mm and at the most 1.5X as long as the gynostegium in M. arizonicum and M. barbigerum) and the more slender shape of the gy- nostegium of M. arizonicum compared to M. bar- bigerum. Metastelma arizonicum and all varieties of M. barbigerum except variety veracruzense have dis- tinctly clavate pollinaria, while those of M. pringlei are ovoid. In M. arizonicum subsp. arizonicum and M. pringlei strongly verrucose trichomes on the co- rolla lobes are absent, while they are present in subspecies chiapense as well as in M. barbigerum. Despite the relatively wide distribution gap be- tween the two subspecies of M. arizonicum (Fig. 3), the morphological differences between the taxa are rather inconspicuous, mainly concerning size dif- Volume 91, Number 1 2004 Liede & Meve 43 Revision of Metastelma ferences and variation in indumentum density on leaves and calyx, as well as the presence of strongly verrucose trichomes on the corolla lobes (present in subsp. chiapense, absent in subsp. arizonicum). Additional specimens examined. EL SALVADOR. Santa Ana: vicin nity of Santa Ana, 28—30 June 1922 (fl). Standley eee (NY). GUATEMALA. Petén: about 3 km NWW of village, 18 Oct. 1961 (fl). 3068 (LL). ы. RAS. El Paraíso: Rio Dantas, 5 July 1962 (fl). 1 did ин. Morazan: entre Las Mesas alme carretera Danlí-Yuscarán. . 1963 (fl). olaa 13162 ( JL. NY); entre km 11- arretera Suyapa a La Montañita, 11 Sep. 1963 (fl). Molina 12851 (NY. ). MEXICO. Chiapas: Cerro del Ocote, ' of Ocozocoautla, ds Oct. 1972 к pa 22 (NY): La Cienega de Leon, 30 km N of Las Cr 30 Nov. 1980 (fr). Breedlove "48083 (MO): Comitán D Dominguez, 21 July 1989 (fl), Liede & Conrad 2 (UBT); Hacienda Monserrate, Purpus 9078 (NY). езш ea: Cerro de Picardo, July 1914 (fl). Purpus 7273 (NY). у Guay nubiles: 5 Linnaea N . Metastelma barbigerum Scheele. 21: 760. 1848. Cynanchum barbigerum (Se a Shinners. Field & Lab. 19: 65. 1951. TYPE: U.S exas: Comal Co., New Braun- fels. jon 1846, E. Lindheimer 459 (lectotype. designated by Blankinship (1907: 143). MO: isotypes, GH not seen, NY!). Figures 4a-1. 5. 2a. Metastelma barbigerum var. barbigerum Plants ascending, 30-80 cm high; shoots annual, glabrous. Leaves with petioles 2-8 mm long. with 0 to 2 colleters at blade base; blades 10—40(—60) х 3—14(-20) mm, ovate, basally rounded, apically acute, or rarely shortly acuminate, adaxially isolat- edly covered mainly on veins and margins with flexuous to recurved trichomes, 250—300 jum long, abaxially glabrous. Inflorescences always one рег node, 2 to 5 chronously, оня ае subsessile to sessile; floral bracts 0.6— . 0.2 mm at the base, ovate; pedicels 4— 6 1 mm bon glabrous. Floral buds 3-3.5 conical to elongated- 2- to 6-flowered, flowers open syn- mm long, 1-1.5 mm diam.. conical: calyx basally fused, rotate to campanulate, abaxially glabrous: calyx lobes 0.8-1.2 X 0.5-0.7 mm, ovate, apically acute; corolla 3.2-5.2 mm long. glabrous, abaxially white to cream (when dry). adaxially white, with slightly verrucose trichomes, 200—500 um long, forming a woolly beard on the apical parts of the lobes and downward-directed hispid stiff trichomes, 300-500 um long. on the central parts of the lobes; lobes fused to ca. % of total corolla length, 0.8—1 mm wide, straight, erect, oblong to lanceolate, apically often rather abruptly recurved, occasionally twisted; narrowing, acute, gynostegial corona 1—1.3 mm long. equaling to lon- ger than the gynostegium; lobes erect to inflexed. attached at the base of the stipe with a distinct hook-shaped base, laminar to solid, subulate. Gy- nostegium 0.7—0.9 mm high, 1-1.2 mm diam., оп a stipe 0.5-0.7 mm long: anthers about as high as broad, trapezoidal, abaxially planar: anther wings 400 jum long. extending along the entire length of the anther, anther wings of adjacent anthers par- allel to each other, centrifugal: connective append- ages 200—300 um lang: 200-300 um wide, equal- ing the slightly inflexed: pollinarium with мерин 400 pm long, be- tween 1.5 times and twice as long as broad, ovoid: caudicles 250 um long, geniculate; pollinia apically attached to the caudicles, ca. 400 um long, 250 um wide, ovate in cross section, cla- vate; style-head white, ca. 0.8 mm diam., 0.5 mm high, upper part 0.25 mm high, equaling the lower part, depressed-conical. Follicles always one per flower, pendulous, 45-70 mm long. 5-6 mm diam. fusiform, terete, apically strongly beaked, medium seeds 5—6 mm long, 3.5— slamen in widt cylindrical, to dark brown, glabrous; 4 mm wide, ovate, medium brown, seta and aseta side smooth, marginally with 0.5-0.6 mm wide wing with entire to slightly lacerate margin: coma 20—25 mm long. Chromosome number. 2n = 22 (voucher: Liede & Mere 2500 (MO, MSUN)) Phenology. Collected in flower from January to October: in fruit from May to November. Distribution. Coahuila, Nuevo San Luis Potosí, Tamaulipas. U.S.A.: . valley bottoms, in Mexico: León. Texas: lime- stone hillsides, shale slopes chaparral, brush or low forest; 30—2300 m. Henrickson (1987: Literature and illustrations. Additional VRAC p MEXICO. Coahuila: Cañon Loma Prieta, m al NE de Hipólito, 23 Oct. 1985 (fl), Villarreal et $ 3115 (LL): Sierra de Parras, 19 Aug. 1982 (fr), Cowan E (NY. TEX). Nuevo León: W of Cerralto, 29 "d 7 (fl). Gregg s.n. (NY): S part of Cerro Pedregoso, 1.5 E V of El Peñuelo, 18 June 1972 (fl), Chiang et al 7964 (LL); Chipinque. 1 July fl), Barkley et al. 5 (TEX); Galeana, ca. 5 km E. of La Poza toward Rio А E 18 Sep. 1933 (fl). Nesom et al. 7592 (LL); Galeana, Ranc ho Aguililla, 16 Sep. 1989 (fl. fr). Hinton et al. 19732 (LL); ca. 120 km 5 of Laredo along wy. 85 to Monterrey, 10 Oct. 1982, Henrickson 19303 (LL); near Monterrey, 2 June 1889 (fl, fr), Pringle 2856 G); Montemorelos, 8 Aug. 1989 (fl), Liede & Conrad 2610 (MO, UBT); 3 mi. W of i er Qu Gentry 6725 ARIZ); Cerca бе Monterrey, 29 June 19 P Fernándes & Barkley 14485 (LL): Monte rey-Laredo ca. 49 km, Sa- linas Victoria, eK 1994. (fl), Hinton et al 24258 (LL): 12.8 mi. S of Sabinas Eo on Hwy. 85, 2 July 1985 fl, fr), Cowan et al. 5359 (LL); tnde iit Bustamante Canyon, 12 Aug. 1988 (A), Patterson 6519 (TEX): San Augustine, Monterrey, 22 Apr. 1960 (fl), Smith M124 (LL). San Luis Potosí: N of the Minas de San Rafael, 30 June 7 44 Annals of the Missouri Botanical Garden . d E r, 1 MAG fi AV КШ ee уута T AR 7 Da, . MON. с H, I DE 7 VEA . Metastelma barbigerum var. r. —d. Corolla а adaxial view. —e, e”. Corona and gynos- tegium. —f. ооа оша —£. Style-head. ae Follic le iy Seed. a, b. : Correll & 1 19695. eg: Liede & Conrad 2610. e': O'Neill 186: s Drawn by J. Conrad. £z — liesnerianum. —i Figure 4. Metastelma barbigerum. a-i. Metastelma bip ibs var. barbigerum. е' “lo abit ud пот rescence, —c. Volume 91, Number 1 2004 Liede & Meve Revision of Metastelma M. barbigerum var. barbigerum M. barbigerum var. liesnerianum M. barbigerum var. veracruzense онь ө М. barbigerum var. barbigerum х М. pringlei ы = — — -115° -110° -105° -100° Figure 5. Distribution of Metastelma barbigerum. 1972 (fl). Chiang et al. 8161D (LL); 15.7 rd. mi. E of El Huizache jet. (at Hwy. 57) on MEX 80 to Cd. del Maiz.. 24 Sep. 1978 (fl). Ж nrickson & Lee еу; is L). Tamau- lipas: 9 mi. NW Chamal. just N of Ojo de Agua. 8 Feb. 1960 (fl). Jio 5063B (LL): Cuatro caminos, Hi- dalgo. 5 June 1994 (fl). Hinton el al. 24186 (LL); ipm 24 Oct. 1978 (f). Calvert 117 (LL): 1 km antes de Pes erca de la Laguna Salada, 22 Se ТД 1985 (fl). а db 2 n 562 (TEX): S of Lomas del Real, 7 mi. N of main Hwy. on dirt road just N of Altamira, 27 Oct. 1959 (fl). Johnston & Graham 4525 (LL): Punta Algodones on N end Y peninsula, 12 Sep. 1981 (fl. fr). Fryxell 3651 (NY. LL): Tampico, 15 June 1897 (fl, fr), Pringle 6646 engin NY): vicinity of Victoria, 1907 (fl). Palmer 360 (NY): : т from Victoria on the road to Casas & Soto la Marina. 3 Oct. 1956 (fl). Martinez & Luyando F-2295 (LL): 5 mi. N of Villagran on the Victoria- Linares Hwy., 12 Nov. 1959 (fr), Johnston & Graham 4674 (LL). State not. known: w northern Mexico, Wright 1676 (NY). U.S.A. Texas: Ban- dera Co.. Rte. 470 betw. Tarpley 2 Unis: at Ио Creek crossing. 16 May 1984 (fl), Ertter 5357 (NY): Bee Co.. 2 mi. SW of Beeville. 17 July 19: EE (l. fr). Correll & Johnston 19695 (NY): Brewster Co.. Big Bend. Oak Springs Trail, 7 Mar. 1989 (fl). Liede & Меге 2 2500 (MO, MSUN); Cameron Co.. dirt road 2 mi. N of Bayview clay bank. 13 June 1967 (fl), Crutchfield 5000 (NY). 5 mi. V of Boc a Chica, 15 Mar. 1966 a, Crute ch el 1125 (NY); tmi. NE of Port Brownsville. SIE June 1967 (fl). S 2984 (NY): El 3 house \ No. 2. along Resaca 3 la Rane ‘ho Vieje . W of Port Isabel. 26 June 1950 d. Webster & Wilbur 3060 (NY): Comal Co.. Old Bear Creek rd. Bear Creek & Little Bear Creek. NW of New Braunfels. 2 fo 1983 N 15 4906 (NY): Duval Co.. S 884. Croft 62 (NY): Gillespie Co.. Balanced Rock, 4 mi. N of Fredericks 1 on TX 965, 23 Aug. 19 78 (ID. Wyatt 1464 (GA); Bear Mountain, 18 Tni 1946 (fl), Correll & Correll 12756 (NY); Kenedy Co.. Yturria Ranch, 6 May ‚ Rocky Bluffs, ede d (NY); Live Oak М о.. пеаг Mathis, 28 June 7 57 (fl), Lundell 8734 (NY); Kimble € 1974 (fl. fr), McGee 134 (GA); Nueces 40, Co.. Christi Bay, Apr. 1894 (fl). Heller 1559 ( along Corpus NY): Corpus Christi, 7 May 1920 (fl). Pennell 10220 (NY): Flour Bluff. 7 Jan. 1939 (fl), Tharp 248 (NY): Pecos Co.. Spring, New Braunfels, July 1851 (fl, fr). Lindheimer 992 (NY): Travis Co., Austin. Eastern Texas, (NY): Barton Springs Creek, 23 July 1943 (fl. fr). Barkley 13360 (NY); Mt. Bonnel. nr. Austin, 24 June 1946 (fl. fr). Albers et al. 46448 (S). County not known: s. coll. (Herb. Berlandianum) 3198 (NY). Wright s.n. 1872, Ha li 5 (NY) Comanche 21 46 Annals of the Missouri Botanical Garden 2b. Metastelma barbigerum Scheele var. lies- О. Williams) Liede & Meve, comb. et stat. nov. Basionym: Cynanchum lies- neriana L. O. Williams, Fieldiana, Bot. 34: 102. 1972. Metastelma pipi (L. O. Williams) Liede, Novon 7: 42. 1997. TYPE: Costa Rica. Guanacaste: vicinty of 48 11 mi. S of Li- beria, 12 Dec. 1969, K. Daubenmire 406 (ho- lotype, F neg. 056988!). Figures 4e', 5 nerianum (L. Plants ascending, 80-200 cm high; subterranean organs constituting a woody rootstock; shoots pe- rennial, basally slightly woody with yellowish bark, glabrous or sparsely covered along a single line with recurved trichomes, 200-250 um long; inter- nodes 2.54 cm long, to 1 mm diam. Leaves with petioles 2-5 mm long, with 0 to 2 colleters at blade blades 20-35 X 4-15 basally obtuse to rounded, apically acute to acu- base; mm, ovate to elliptic, minate, adaxially glabrous or with isolated recurved trichomes, 200—300 um long mainly on veins and margins, abaxially glabrous, Inflorescences always one per node, 2- to 5-flowered, 1 to 5 flowers open synchronously, sciadioidal, bostrychoid, sessile to pedunculate with to 3 mm long peduncles, glabrous or sparsely covered along a single line with re- curved trichomes, 150-200 um long; rachis 1—2 mm long, straight; floral bracts 4—6 X 2-3 mm at the base, deltoid to ovate, glabrous; pedicels 2—3 mm long, glabrous. Floral buds 2-2.5 mm long, 1— 1.3 mm diam., ovoid to conical; calyx basally fused, rotate to campanulate, abaxially with a few tri- chomes on the calyx tube (characteristic for this 0.2-0.4 mm, apically obtuse to acute; corolla 2-3 mm long, variety); calyx lobes 0.7-1 х ovale, abaxially white, glabrous, adaxially white, with ver- rucose trichomes, 100—150 um long, on the apical and on the lateral parts of the lobes, and smooth, downward-directed trichomes, 250-300 um long, on the central parts of the lobes: lobes 0,8—1 mm wide, suberect, apically acute, recurved, with straight to slightly revolute margins: gynostegial co- rona 1—1.2 mm long. equaling to slightly longer than the gynostegium: lobes erect, attached near the base of the stipe, laminar. ovate, with entire margins. Gynostegium 0.4-0.6 mm high, 0.8-1 mm diam., on a stipe 0.3-0.4 mm long: anthers broader than high, trapezoidal, abaxially planar to convex: anther wings 300—400 jum long, extending along the entire length of the anther, anther wings of ad- jacent anthers parallel to each other, in the same plane as the anther to slightly centrifugal: connec- tive appendages 300-400 um long, 200—300 um wide, ovate to triangular, equaling the stamen in width, slightly inflexed; pollinarium with corpus- culum 150-200 um long, between 1.5 times and twice as long as broad, ovoid; caudicles 60—80 jum long, cylindrical to flattened, s-shaped, concave- convex; pollinia apically attached to the caudicles, 150-200 um long, 80—100 um wide, clavate; style- head white, ca. 0.5 mm diam., ca. 0.3 mm high. upper part ca. 0.15 mm high, equaling to longer than the lower part, flat to depressed-conical. Fol- licles always one per flower, pendulous, 35—40 mm long, 2 mm diam., terete, apically strongly beaked, dark brown when dried, glabrous. Phenology. Collected in flower from January to August and in October; in fruit in January. Distribution. Belize; Costa Rica: Guanacaste: Guatemala: Péten; Mexico: Campeche, Chiapas, Ta- basco, Yucatán; close to rivers, on disturbed slopes; low altitudes, to 200 m. Literature апа illustrations. 103, fig. 1A Williams (1972: Additional оү — BELIZE. Hectar ; ver, 6 Маг. 1935 (fl), Gentle 1508 (NY); ‚12 ka N of Belus. pa Aug. 1936 (fl), O'Neill WIS): Mile 5 % Northern Hwy., 7 June 1974, Dwyer 12774 (NY): Vaca, 25 Apr. 1938 (fl), Dele 2517 (NY); betw. Belmopan & 2 Ignacio, 13 Feb. 5 (fl), Liede 3242 (ОВТ). GUATEMALA. Petén: я Lagunas, El Cedro, 4 Мау 1969 ^" fr), Contreras 8466 (NY, TEX); Tikal Nat. 79 7050 ded de Santa Fe, 29 Jan. 1960, Contreras 532 (L " ). Campeche: 2 Oct. 1847 (fl) wards s.r de Й = al Sureste de Conhuas dirección Calcul. 150 m, 4 Aug. 1995 (fl), Gutiérrez 4632 (TEX). e oo ate Palenque, 16 July й (fl), Машаа 3765 (LL ; 17 km N of Tuxtla Gutiérrez се road to El Nees 30 Oct. 1965 Le Breedlove 14010 0(T Tabasco: ае 8 July 1962 (Е 31376 (WIS 28/17 (WIS) Yucatan: lzamal, UPS, WI Y. 2 = ® ч a> Cs 1895, Gaumer 539 (NY 2c. Metastelma barbigerum Scheele var. vera- cruzense Liede & Meve, var. nov. TYPE: Mexico. Veracruz: 48 km SE of Nautla, 30 Dec. 1972, J. Taylor & C. Taylor 12532 (ho- lotype, NY!). Figure 5. Haec varietas varietati liesneriano similis, sed differt pedicellis calycibusque pubescentibus. Plants ascending, 200 cm high; shoots perennial, covered along a single, but occasionally a broad line of recurved to flexuous trichomes, 300—400 jum long. Leaves with petioles 2-7 mm long, with O to blades 10-45 X 6-15 mm, basally obtuse to rounded, apically acuminate 2 colleters at blade base; to acute, adaxially glabrous or with isolated re- curved trichomes, 250-300 um long. mainly on veins and margins; abaxially sparsely covered only on the midrib with recurved trichomes, ca. 300 jum Volume 91, Number 1 2004 Liede & Meve 47 Revision of Metastelma long. Inflorescences always one per node, 2- to 7- flowered, 2 to 7 flowers open synchronously, scia- dioidal: peduncles 1—4 mm long. sparsely to densely covered along a single line or over the entire sur- face with flexuous trichomes, 150-200 um long: floral bracts 040.8 X 0.2-0.3 mm at the base, triangular to ovate, sparsely with trichomes on the surface: pedicels 2-4 mm long. densely covered over the entire surface with flexuous trichomes, ca. 200 um long. Floral buds ca. 2 mm long. ca. | mm diam.. conical, with imbricate aestivation: calyx ba- sally fused. rotate to campanulate, ciliate or abax- ially with trichomes (mainly on the tube): calyx lobes 1—1.2 X 0.5-0.6 mm, ovate, apically acute: corolla rotate to campanulate, 2-3 mm long. abax- ially white, glabrous, adaxially white, adaxially with verrucose trichomes, 100—150 um long. on the api- cal and on the lateral parts of the lobes and down- 250-300 um long. on the central parts of the lobes: lobes fused ward-directed smooth trichomes, to ca. % of total corolla length, 0.6-0.8 mm wide. erect to patent, ovate, apically acute: gynostegial corona 0.8—1 mm long, equaling to longer than the eynostegium: lobes erect, attached along the stipe. laminar, very narrowly ovate. Gynostegium ca. 0.6 mm high, са. | mm diam., on a stipe 0.3-0.5 mm long: anthers broader than high, trapezoidal, abax- ially planar: anther wings 300 um long. extending along the entire length of the anther, anther wings of adjacent anthers parallel to each other. in the same plane as the anther: connective appendages 300—350 um long, 250-300 um wide, ovate, nar- rower than the stamen, slightly inflexed; pollinar- ium with corpusculum 150-200 jum long. between 1.5 times and twice as long as broad, elliptic; cau- dicles 50-60 um long. flattened, s-shaped, con- cave-convex; pollinia subapically attached to the i 200 um long, ca. caudicles, ca. 100 pm wide, ovoid: style-head white. ca. 0.7 mm diam ca. 0.5 mm high. upper part ca. 0.25 mm high. equaling the lower part. flat to depressed-conical. Follicles always one per flower. 30-40 mm long. 4-6 mm diam., obclavate, straight, apically strongly beaked, green-brown, glabrous: seeds 5-6 mm long. 2.5—3 mm wide, ovate, medium to dark brown, seta side tuberculate, papillose with regularly arranged pa- pillae, aseta side vaguely tuberculate, marginally with 0.2-0.3 mm wide wing with denticulate mar- gin; coma 25—30 mm long. Phenology. | Collected in flower from August to December: in fruit in December. Distribution. Mexico: Tabasco. Veracruz: sand dunes and savanna borders; 0—50 m. Cowan & Magafia 3270 (NY) from Huimanguil- lo. Tabasco. is not entirely. glabrous like variety liesnerianum, but not as densely indumented as the other Metastelma barbigerum var. veracruzense specimens. There is a second NY specimen of Purpus 2094. which is referred to M. schlechtendalit var. schle- chtendalit. The Metastelma barbigerum complex is charac- terized by mostly glabrous stems (but with a single line of trichomes in var. veracuzense), conically shaped gynostegia with obliquely protruding anther wings (guide rails) and corona lobes that hardly ex- ceed the anthers (Figs. la. c. e. f: compare also 1987). fer in the attachment of the corona lobes, which is Henrickson, The two southern varieties dif- not as distinctly hook-shaped as in the typical va- riety. The distribution of this complex is confined to the eastern landmasses surrounding the Gulf of Mexico. Tabasco: Huimanguillo, km 39.7 de la desviación de Humanguillo hacia Fco. 10 Oct. 1980 (f). Cowan & Magaña 3270 (NY). Vera- eruz: hte. 180, 130 IW Catamaca. 16 Sep. 1970 (fl). Burch 2984 (NY): rw pid s de la planta Laguna Verde. 11 vw. 1975 (fl), Dorantes 5270 (NY): about 4.5 mi. W of Palmilla along pui through Huatusco, 10 Aug. 1971 (fl). Stevens 1409 ( : Tecolutla. 20 Dee. 1963 (fl. fr). Cavender s.n. (WIS): zi uapan and vicinity, Nov. 1906 (fl). Purpus 2094 (HBG, NY). Paratypes. MEXICO Rueda, 3. Metastelma californicum Benth.. in Hinds. R. B., Bot. Voy. Sulphur: 33, pl. 18. 1844. Cy- nanchum californicum (Benth.) Moran, in Lev- in & Moran, Vascular Fl. Isla Socorro Mexico (Mem. San Diego Soc. Nat. Hist. 16): 26. 1989. TYPE: Mexico. Baja California: Magdalena Bay, С. Bentham s.n. (holotype. K!). 6a-f. Figures За. Metastelma californicum Benth. subsp. eal- ifornicum Plants ascending, 50-150 em high: shoots an- nual, basally woody, with gray-brown bark, glabrous or isolatedly glabrescent along a single line with flexuous trichomes, 200—300 um long: internodes 2—4 cm long, 0.5-0.8 mm diam. Leaves with peti- oles 3-6 mm long. with 2 colleters at blade base: blades 10-20(-25) х 4—10(-14) mm. ovate to el- liptic, basally rounded, apically acute to acuminate. adaxially sparsely covered on the midrib and the 200—300 um long, abaxially glabrous. Inflorescences always one 8 ys l › margins with flexuous trichomes, per node, 2- to 6-flowered, 2 to 3 flowers open syn- chronously, sciadioidal, floral bracts 0.5—0.6 X 0.2-0.3 mm at the base, triangular, gla- subsessile: 48 Annals of the Missouri Botanical Garden „Kéi Amie 409 as Wi WS E ASA Wr \ . . 7 y me {, И М, E Ng A - \ i 1 mm Figure 6. усун Ima californicum subsp. californicum. lobe, adaxial view. —d. Gynostegium and corona. —e Conrad 2955. us by J. Conrac brous; pedicels 3—4 mm long, glabrous or with a few isolated trichomes. Floral buds ca. 2 mm long. ca. 1.5 mm diam., ovoid; calyx basally fused, campan- agn vu glandular, glabrous: calyx lobes 1— ).4—0.6 mm, ovate, apically acute; corolla pan Из to rotate, 2.5—3.5 mm long, abaxially glabrous, 200-250 white, purplish along the main nerves. adaxially white, with verrucose trichomes, um long. on the apical and on the lateral parts of the lobes and with smooth, downward-directed tri- chomes, 180-220 jum long, on the central parts of . Shoot и inflorescences. —b. Flower. —c. Corolla Style-head. a: Brandegee s.n. b-f: Liede & Pollinarium. the lobes: lobes basally fused, 1-1.2 mm wide, pat- ent to erect, ovate, apically acute to obtuse; gynos- tegial corona 1.3-1.5 mm long, slightly longer than the gynostegium; lobes erect, attached directly un- lanceolate. Gynos- derneath the anthers, laminar, tegium ca. | mm high, ca. 0.9 mm diam., sessile: anthers higher than broad, rectangular, abaxially gibbous; anther wings ca. 500 рт long, extending along the entire length of the anther or beyond, anther wings of adjacent anthers divergent toward the base, in the same plane as the anther; connec- Volume 91, Number 1 2004 Liede & Meve Revision of Metastelma tive appendages 300-400 um long. ca. 300 рт wide, ovate, equaling the stamen in width. slightly inflexed: pollinarium with corpusculum. 140—150 um long. between 1.5 times and twice as long as broad, ovoid: caudicles 60-70 рт long, cylindrical to flattened, straight. ascending, slightly thickened at the insertion of the pollinium: pollinia subapi- cally attached to the caudicles, 180-220 jum long. 80-90 um wide, ovate in cross section, ovoid: style- head white. ca. 0.5 mm diam., ca. 0.4 mm high, upper part ca. 0.2 mm high. equaling the lower vart, conical. Follicles always one per flower, 50— 60 mm long, 4-6 mm diam., fusiform, terete. api- cally strongly beaked. wingless, dark brown. gla- brous; seeds 5-6.5 mm long. 2.5-3 mm wide, ovate, dark brown, seta and aseta side tuberculate, mar- ginally wingless, entire: coma 10-20 mm long. Phenology. Collected in flower from October to April and in July and August: in fruit in December, February, March. and October. Distribution. Mexico: Baja California, Sinaloa, Sonora, Revillagigedo Islands: on rocky slopes and in sandy washes, very dry desert. vegetation; 0— 6 m. Literature and illustrations. Bentham (1844: pl. 18). Additional specimens | examined. MEXICO. Baja Cal- ifornia: 3.2 mi. NW of Arroyo Palmarito via Hwy. 1. 21 July 1990 (fl. fn. — & Van Devender 7661 (ARIZ): E co. 21 Mar. ү (fl. fr). Shreve 7197 » of Arroyo Sec (ARIZ): Catavina, 76 mi. El Rosario on Mex. Hwy. 1, 13 Dec. 1977, Nixon & n 831 (LL); 2 mi. NW of Catavina, 29 Apr. 1980 (fl). Davis & ADR 66297 (В); Cerro Tordillo & vicinity, Systema de la Sierra Vis- caino, 12 Mar. 1947 (Н), Gentry 7438 (ARIZ): Escondito Creek, 7 Mar. 1934 (fl. fr), Shreve pons (ARIZ): Escon- dido Canvon betw. Punta Prieta M ocean, 7 Mar. 193 | (fl) Ferris 8589 (NY): 8 mi. N of Mt. Mesquital Grande, 9 Feb. 1935 (fl), Haynes & Stewart s.n. (NY); O mi. S of Laguna Chapala к а. 17 Oct. 1959 (f, Wiggins 15048 (ARIZ); Picacho de Santa Clara, Nov. 1947 (fl), Gentry 7716 (ARIZ): Playa S бай Pedrito. Los Cerritos, al SW de Todos Santos, 13 Oct. 1985 (fl. fr). Tenorio et al. 10441 (MO); 10-20 km N of Puerto Santa Catarina on rd. to Rancho Santa Catarina, ] ir 1985 (fD. Breedlove 62261 (NY); 20-30 mi. 5 of Punta Canoas along da to Punta Cono, 27 Feb. 1986 (fl). ا‎ 62528 (LL); 12.1 mi. (by rd.) N of Pi i 3 we 17 Oct. 1967 (fl). x wd & Turner 67-64 (ARIZ): 2.5 mi. 5 of 9 9 Prieta, 17 Oct. 1959 (fl. A Wiggins 15085 (ARIZ); just 5 of Rosar- ito, ^ Mar. 1979 (fl. fr). 5 & Bowers 2119 (ARIZ); Todos Santos, ca. 8 km on track to Sierra 1 aguna, 20 ppa 1992 (fl. fr). Liede & Conrad 2955 (l LM): Todos sete ys S.H. S ae Inlet. Magdalena Bay. 10 Aug. (fl), Howell 10639 (NY). Sinaloa: Isla Tac hec hilte. : en m. 20 Jan. 1945. Gentry 7136 (ARIZ NY). E sla San уе Nolasco, E side of ad, Nov. . Felger 9635 ане 5 Li Las los ‘as on the coast = de Gulf of California. ‘eb. 1992 (fl. fr), Van Devender et al. 92-121 (ARIZ): ps Santos, Jan. 1890 (fl). 1995, C homajabire s, 1.8 mi. N of Huatabampito, 23 Dec. Revil- Fishbein & Mc Mahon 2750 (ARIZ), 2769 (ARIZ). lagigedo Islands: Socorro Island. 24 May 1903, Barkelew 217 (ARIZ, NY). 3b. Metastelma californicum Benth. subsp. lanceolatum (Schltr.) Liede & Meve. comb. Basionym: Metastelma lanceola- Bull. Herb. Boissier 6: 840. 1900. San José Pass. et stat. tum Schltr., TYPE: Mexico. San Luis Potosí: $ 12 July 1890, C. G. Pringle 3138 A M!: HBG!, NY! nov. designated here, isotypes, Figures 7a-g. 8. Ditassa mexicana Brandegee. Calif. Publ. Bot. 10: 414. 1924. Syn. nov. T cienda Montserrate, Sep. 1923, ( (holotype, UC 220412 not seen: не GH! NY!) Univ. "PE Plants ascending. to 250 cm high: shoots peren- nial. basally woody, with brownish bark. sparsely covered along a single line with recurved tri- chomes, 150—300 рт long: internodes 1.7 cm long. to 1 mm diam. Leaves with petioles 3—5 mm long. with 2 colleters at blade base: blades 15-35 X 3-16 mm, ovate to oblong, basally rounded, api- cally acute to acuminate, adaxially sparsely cov- ered only on the midrib and the margins or over the entire surface (Roe & Roe 1885) with erect to recurved trichomes, 200—350 рт long. abaxially glabrous (occasionally with a few trichomes on the midrib). Inflorescences always one per node, 4- to 7-flowered, 1 to 7 flowers open synchronously, scia- dioidal, subsessile; floral bracts 0.5-0.6 X 0.3-04 mm at the base, triangular, glabrous or ciliate: ped- icels 2-3 mm long. glabrous or sparsely covered along a single line with flexuous trichomes. ca. 200 um long (Roe & Roe 1885). Floral buds 2-2.5 mm 1.3-1.5 mm diam., ovoid: calyx basally fused, campanulate, abaxially glabrous or with trichomes on the tube (Roe & Roe 1885): calyx lobes 0.8—1 X 0.4-0.6 mm, ovate, apically acute; corolla rotate long, to eyathiform, 2.5—3 mm long. abaxially greenish adaxially white or cream, with verrucose tri- chomes, 120-180 um long, on the apical and on the lateral parts of the lobes and smooth trichomes. 180—200 um long, on the central parts of the lobes: 1—1.2 mm wide, patent. ovate. white, lobes basally fused, apically acute to obtuse: gynostegial corona 0.6— 1.2 mm long. equaling to longer than the gynoste- gium; lobes erect, attached directly underneath the anthers, laminar, ovate, apically acute to longly acuminate. Gynostegium 1-1.3 mm high. 1.2-1.5 mm diam., sessile: anthers about as high as broad. anther wings 500— rectangular, abaxially gibbous: 600 um long. extending along the entire length of the anther, anther wings of adjacent anthers par- Annals of the Missouri Botanical Garden ж ä (Lua LET! S He К * = V bay ^ Ve үз, NS NX NE E "EC EN We m t Ам N М, mE Figure 7. Metastelma californic um шк lanceolatum. —а. Habit. —b. Shoot with inflorescence. —c, c'. Flower. — d. ( ‘orolla lobe, adaxia view. —e, e' na and gynostegium. —f. Pollinarium. —g. S drawn by J. Conrad. e”, e”. Roe & Roe 1885. executed by U. x. Style-head. a-g: Purpus 9167, Meve after sketch by J. Conrad Volume 91, Number 1 Liede & Meve Revision of Metastelma 2004 ГЕЯ М. wi T , $ ed س‎ | |p|] AA | | 2); | A | $ E Lem 88. — ve | — 7 NM 35? f À | | | | | اہ‎ ~ A, 14 | . | : V і i os: | | | X — M \ | | |S „Л daa o o. i == Х - X crap ГТ" 30 " * 1 iz * N уу | ea S "uii s 4 . n Б | | A | p e. 4 t X. y ‘ | Б 4\ etj AS le | a A ‚ J | - AAA AAA : 25° e ^ ] | ү у ү 44 i | ] PY y | Te \ Е =y Ж 4 — 4 1 ыч i= 2 > 4 20° | hs de ROS | А | e | i > А + ; = к< 07 | | "nM "a 4 4 44 s — | N californicum ssp. californicum : | Т F | | | А M.californicum ssp. lanceolatum 4 i | | -120° -115° -110° -105° -100° -95° -90° Figure 8. Distribution of Metastelma californicum. allel to each other, in the same plane as the anther: 1. 400 connective appendages ca. 400 um long. cz width, pm wide, ovate, equaling the stamen in slightly inflexed: pollinarium with corpusculum 160-180 um long. between 1.5 times and twice as long as broad, ovoid; caudicles 60-80 um long. cylindrical to flattened, convexly recurved; pollinia subapically attached to the caudicles, 280-300 um long, са. 120 рт wide, ovate in cross section, ovoid: style-head white, 0.7— 0.4—0.6 mm high. upper part 0.3— longer than the lower + horizontal to straight, 0.9 mm diam.. 0.4 mm high. equaling part. conical. Follicles always one per flower, 35— 45 mm long. 4-6 mm diam., obclavate. terete, api- cally strongly beaked, dark brown. longitudinally grooved: seeds ca. 5 mm long. ca. 3 mm wide. ovate, medium brown. seta and aseta side smooth, mar- ginally with 2 mm wide wing with denticulate mar- gin: coma 15-25 mm long. Phenology. Collected in flower in February. May. and from July to October: in fruit in February, August, and September. Distribution. Mexico: Chiapas, Guerrero, Hi- dalgo. Jalisco. México Distrito Féderal, Morelos. Oaxaca, Puebla, San Luis Potosí. Yucatán: on lime- stone slopes in scrub vegetation and deciduous tropical forest: 1000-2200 m. For Metastelma lanceolatum. cited the type as follows: pes prope Zoquitlan, flor. Jun. C. Seler & H. Seler ° However. the first set of collections of Seler n бе was housed in B (Vegter, 1986), and the type of M. lanceolatum was probably destroyed during the second World War. It is not duplicated in any of the herbaria consulted for this study. The specimen selected as neotype (Pringle 5136. NY) “Metastelma lanceolatum Schlechter (1906) “Mexico. Oaxaca: Distr. bears the annotation Schltr.” in Schlechters own handwriting. The protologue of Ditassa mexicana states. ex- plicitly “corona duplici." which is the generic char- acter of Ditassa. No other "Ditassa^ specimen ex- cept for the type of Ditassa mexicana was ever collected in Central America. However. none of the flowers examined had a double corona. This spec- imen exhibits the characters of Metastelma califor- nicum subsp. lanceolatum. so that Ditassa remains a genus restricted to South America. Metastelma californicum is easily distinguished Annals of the Missouri Botanical Garden from all other Central American Metastelma species by a stout, sessile gynostegium with anther wings (guide rails) as long as the gynostegium. The corona is spreading and hardly exceeds the gynostegium. Morphological differences between the two subspe- cies of M. californicum mainly concern the shape of the corona lobes. As separation is both geograph- ical and ecological, with the typical subspecies oc- curring in very dry desert vegetation below 640 m, and M. californicum subsp. lanceolatum in scrub vegetation and deciduous tropical forest between 1000 and 2200 m, subspecific rank, as in the par- allel case of M. arizonicum, is warranted. — онн 5 ME XICO. sei Federal: n Hwy l i "Roe 1885 (NY. WIS); Volcán, 2 Feb. 1935 (fl, fr), Hinton et al. 7310 (NY). Guerrero: 10 m Copac Cp 5 Aug. 1970 (fl), Boege 1438 R); La Posa del Lago Tres dd чорва abana, 19 Sep. pus (fl). Boege à (FR) I lidal donde 'a de Tolan- tongo, 8 Aug. 1991 (fl). id „ МҮ); slopes of Barranca de шл 8 Aug. 1948, n & Wood 4409 (LL); El Capulin, near km 1 ^ on highway be & пага 4 Aug. 1948 (LL); Jacala, 2 July 1939 Ny Chase 7418 (ARIZ, NY). L). Me 9 (fl), Pringle 10838 (ARIZ, TEX). Oa- idein Galeotti 1571 (NY). Puebla: 11 km al SE de San Juan Ixc ке 5 du 1986 (fl), Salinas & Tenorio 3294 (ARIZ, LL). San Luis Potosí: Pedro, 29 July 1934 (fl). edm 17729 (NY); . 1939 (fl, fr), Shreve 9312 (ARIZ); Hwy. Valles to 25 May 1981 (fl), Fry xell & An- de 7 (NY). Yucatán: above Calcehtok, 21 Feb. 1982 (fl), White & Mott 74 (NY). 4. Metastelma cuneatum Brandegee, Zoe 5: 216. 1905. TYPE: Mexico. Sinaloa: Yerba Buena, vicinity of Culiacan, 10 Sep. 1904, T. S. Brandegee s.n. (holotype, UC not seen; iso- types, GH!, US). Figures 9a—h, Plants ascending, 200—300 cm high; shoots pe- rennial, basally woody with grayish brown bark, sparsely to densely covered along a single line with erect to recurved trichomes, 300—400 рт long; in- ternodes 1.8-3 cm long, ca. 1 mm diam. Leaves with petioles 2—4 mm long, with 2 colleters at blade base; blades 15-20 х basally obtuse, apically obtuse or acuminate, adax- 3-7 mm, ovate to oblong, ially sparsely covered, only on the midrib with ap- pressed to recurved trichomes, 200-300 um long, abaxially glabrous. Inflorescences one, occasionally two per node, 3- to 8-flowered, 2 to 6 flowers open synchronously, sciadioidal; peduncles 0.5-3 mm long, sparsely covered along a single line with erect trichomes, 150-200 um long; floral bracts 0.2—0.5 ).2-0.3 mm at the base, triangular, elongated- triangular or ovate, with scattered trichomes on the surface; pedicels 3-4 mm long, glabrous, isolatedly to sparsely covered along a single line with flexuous trichomes, 200-250 um long. Floral buds 1-1.3 mm long, 0.8-1 mm diam.; of its length, campanulate, abaxially glandular; ca- lyx lobes 0.6-0.8 X acute; corolla rotate, 1.5-2 mm long, abaxially yel- lowish green, glabrous, adaxially white, with ver- calyx fused for ca. Y .2-0.3 mm, ovate, apically rucose trichomes, 50-100 рт long, on the apical and on the lateral parts of the lobes; and with a few slightly longer, retrorse and smooth trichomes on the central parts of the lobes; lobes basally fused, 0.3-0.5 mm wide, erect to incurved, ovate, apically acute; gynostegial corona 0.5—0.7 mm long, equal- ing to shorter than the gynostegium: lobes erect to inflexed, attached directly underneath the anthers, laminar, ovate to triangular. Gynostegium 0.5-0.6 mm high, 0.7—0.8 mm diam., sessile to on a column ca. 0.3 mm long: anthers about as high as broad, rectangular, abaxially convex; anther wings 500 um long, extending along the entire length of the anther, anther wings of adjacent anthers par- allel to each other, basally widened, in the same plane as the anther, basally forming a distinct "mouth" with the basal lateral margin of the anther; connective appendages ca. 500 jum long, ca. 400 pm wide, ovate, equaling the stamen in width, strongly inflexed; pollinarium with corpusculum ca. 120 um long, more than twice as long as broad, slenderly oblongoid; caudicles ca. 100 um long, cylindrical, s-shaped, convex-concave, thickened at the insertion of the pollinium; pollinia subapically attached to the caudicles, ca. 150 um long, ca. 50 um wide, ovate in cross section, reniform; style- head ca. 0.7 mm diam., 0.35 mm high, forming distinct noses at upper ends of corpusculi, upper part ca. 0.15 mm high, shorter than the lower part, depressed-conical. Follicles always one per flower, 60—80 mm long, 4—5 mm diam., siform, very slender fu- terete, apically shortly beaked, grayish brown, glabrous; seeds 6-7 mm long, 3-3.5 mm wide, pyriform, medium brown, seta side vaguely tuberculate, aseta side smooth, marginally with 0.2-0.3 mm wide wing with denticulate margin; coma 25—30 mm long. Phenology. Collected in flower in September and from November to January; in fruit in January. Distribution. Mexico: and Sinaloan thornscrub; 0-25 m The inconspicuous indumentum of the corolla, Sinaloa, Sonora; coastal the anther wings forming a distinct “mouth,” and the very slenderly oblongoid corpusculi carrying Volume 91, Number 1 Liede & Meve 53 4 Revision of Metastelma с. Flower. —d. Corolla lobe. adaxial Figure O. Metastelma cuneatum. а. Shoot with follicle. —b'. Inflorescence. o view. —e. Gynostegium and corona. —f. Pollinarium. —g. Style-head. —h”. Seed. a. eg: Friedman 150-95. b: van Devender et al. 1284. h": Friedman 199-95. Executed by U. Meve after sketch by J. Conrad. Annals of the Missouri Botanical Garden O Metastelma cuneatum А Metastelma latifolium | Ф Metastelma minutiflorum 30° — -> A A 25° -115° -110° Figure 10. markedly s-shaped caudicles make this coastal res- ident an easily recognizable species. Additional specimens examined. in arroyo d оа, 1.5 K MEXIC Sinaloa: m NE of Cana, 15 m, 17 Jan. 19¢ 5 (fl), Frie Ed 015-95 (ARIZ); е Muerto, 1.5 Т. NNW of Ca ›аһштоа on rd. to С Bo 17 Jan. 1995 (fl), Feldman. 104-95 (ARIZ); 9.8 km W and 5 km N of Mex 15 on Las Bon 'as Rd., 20 Jan. 1995 fr), Friedman 150-95 (ARIZ): 6.25 km E of Camahui- roa on rd. to Diez de Abril, 20 Jan. 1995 dl, fr), Friedman 199-95 (ARIZ): 4.4 km SW Ejido Tierra y Libertad. 1 km Von dirt rd., 19 Sep. 1994 a. Friedman rines 94 (ARIZ). Sonora: along rd. to Bachoco, ca. 3.5 mi. 24 Dec. 1995 (fl), Fishbein 27. 59 (ARIZ the coast of the Gulf of California, 1 Feb. 1992 (fl, fr), Van Devender et al. 92-133 (ARIZ); 1.5 km NNW of Ca- mahuiroa on rd. to Las Bocas, 23 Nov. 1993 (fl. fr). Van Devender 93-1253A (ARIZ); 2 km W of Tierra y Libertad on northern rd. to nU ya, 24 Nov. 1993 (fl), 150 Devender et al. 93-1284 (ARIZ); mouth of arroyo Masiacz Las Bocas on the Gulf of California, 22 Sep. 1994 (fl), Van Devender & Yetman 94-698 (ARIZ). — E of Las Bocas, ); Las Bocas on 20° -105° -100° Distribution of Metastelma cuneatum. M. latifolium, and М. minutiflorum. э. Metastelma eliasianum Dugand, Caldasia 9: 445. 1966. TYPE: Colombia. Bolivar: N of Ar- jona, 30-50 m, 15 Nov. 1926, E. P. Killip А. C. Smith 14531 (holotype, US 1.350.518 not seen; isotypes, GH not seen, NY!, P!). Fig- ures lla-f, 12 C ynanchum infimicola L. O. Williams, Ann. Missouri Bot. "ard. 55: 48. 1968. Syn. nov. Metastelma о (L. О. о W. D. Stevens, 5 1988. TYPE: = "s ere K-2 го; 1967, J. D. r & S. | Hayden 7. 7540 Гата type, F!; pl MO not abe : 00). Plants ascending: shoots perennial, basally woody, with yellowish to brownish bark, sparsely covered along a single line with recurved tri- chomes, 150-250 um long; internodes 3-7 cm ong, 1-1.5 mm diam. Leaves with petioles 6—10 mm long, with 2 or 3 colleters at blade base; blades 25—50 X 8—30 mm, ovate, basally rounded to cu- Volume 91, Number 1 Liede & Meve 55 Revision of Metastelma ois! 7 go £x Е \ TE \ Figure 11. d. Gynostegium and corona. Drawn by J. Conrad. . Pollinarium. H. neale, apically very acute, adaxially isolatedly cov- ered only on the midrib with appressed trichomes, ca. 250 um long, abaxially glabrous. Inflorescences always one per node, 4- to 7-flowered, 2 to 4 flowers open synchronously, bostrychoid; peduncles 5-15 mm long, sparsely to densely covered along a single line with recurved trichomes, 200-250 рт long; rachis 0-5 mm long, angular; floral bracts 4—5 X 3—4 mm at the base, ovate, papillose: pedicels 2-5 Metastelma para —a. Shoot des inflorescences. —b. F Style-head. a: Ие KES ES — > 3 3 A AER: e "PI Corolla lobe, adaxial view. — Allen & Seibert 1499. bf: Duke 12414. "lower. —¢ mm long, glabrous. Floral buds ca. 1.5 mm long. ca. 1 mm diam., cylindrical; calyx basally fused, cyathiform, abaxially glabrous or with a few sparse trichomes on the tube; calyx lobes 0.8-1 х 0.4— 0.5 mm, ovate, apically acute; corolla campanulate, 2-2.5 adaxially white, with verrucose, 120-150 um long trichomes on the apical parts of the lobes: lobes fused for ca. % of total corolla length, ca. 1.5 mm mm long, abaxially greenish white. glabrous. Annals of the Missouri Botanical Garden o | | p | Vas 4 X 25 | %: AL | — Y ALO | P | Й — — NS -x n | vM е — 7 OW | о M | ' = \ "E ui | 1 Du ы x | 7 | WE. В ` 4 — — — — 20° f | сә qc 7 * ~ м | p | N / | 8 "n pm id EI ^A Bac — гч مي‎ ~ А y — ا‎ | > + 4 E: "ed -—* AR, - "rs Ua _> : | v2 M. | TU AU DNE: “eens 2 т | 15° : A 5 д у“ | З P al : [. 7 | № * 1 \ | — AS А: А | o 3 | | A 7 et r Xie N P m i | X r Р, А \ 1 | -— X | le 10° 1081 М Чад, > ls ) e Metastelma eliasianum ^m E — ^ кы P > ae N А Metastelma pedunculare * „% T XA N ш Metastelma stenomeres QA) 1 3 ү 2 — ae | | ل‎ — - 5° -100° -95° -90° -85° -80° -75° Figure 12. Distribution of Metastelma eliasianum, M. pedunculare, and M. stenomeres. wide, patent, oblong, apically acute; gynostegial co- Phenology. Collected in flower in May and rona 1-1.2 mm long, equaling the gynostegium; lobes erect, apically reflexed, attached at the base of the stipe, laminar, oblong. Gynostegium ca. 0.3 mm high, ca. 0.8 mm diam., on a stipe 0.8-1 mm long; anthers broader than high, rectangular, abax- ially planar; anther wings 300 um long, extending along the entire length of the anther, anther wings of adjacent anthers parallel to each other, in the same plane as the anther; connective appendages ca. 150 um long, ca. 300 um wide, ovate, equaling the stamen in width, strongly inflexed; pollinarium with corpusculum 100 um long, about twice as long as broad, margins of the corpuscular cleft parallel: caudicles 60-80 jum long, cylindrical, s-shaped, convex-concave; pollinia apically attached to the caudicles, ca. 350 jum long. ca. 70 jum wide, ovate in cross section, clavate; style-head white, ca. 0.7 mm diam., ca. 0.2 mm high, flat. Follicles one per flower, 50—60 mm long, 3—4 mm diam.. obclavate tc narrowly fusiform, terete, apically strongly = beaked, wingless, medium brown, glabrous; seeds unknown. from July to September. Distribution. Colombia. Panama: Coclé, Canal Zone, Panamá; savanna and thickets; 0-200 m. one 2, NY) from 700 m. Metastelma E is most closely related to record (Tyson et al. 3962 M. pedunculare and shares with it the long stipe, the tiny, ca. 0.3 mm high gynostegium, and a sim- ilar pollinarium structure. However, the long corona lobes of M. eliasianum originate at the base of the stipe, while the shorter corona lobes of M. pedun- culare originate just underneath the gynostegium. Metastelma eliasianum thus takes an intermediate position both in corona morphology and geograph- ical distribution between the truly Central Ameri- can M. pedunculare and the South American M. parviflorum, in which the attachment of the corona lobes has moved even further away from the gy- nostegium and onto the base of the corolla. Based on specimen records, Metastelma eliasian- um might be more widely distributed in South America, but this is beyond the scope of the present revision. Volume 91, Number 1 2004 Liede & Meve 57 Revision of Metastelma б PANAMA. Coclé betw. Las Margeritas & El Va d 1938 (fl), Won. 65 (NY). Panamá: E mi. S of El Valle May 1966 (fl), Tyson et al. 3962 (NY); nr. Rio Mar, Е July 1967 (Н), Duke 12414 (NY. WIS): Taboga Island. Sep. 1962 (fl), Dwyer 2809 (NY), 20 May 1911 (fl), а 3539 (NY), 23 July 1938 (fl). Woodson et al. 1499 (NY). Additional specimens T 6. Metastelma latifolium Rose, Contr. U.S. Natl. Herb. 1: 106. 1891. TYPE: Mexico. Sonora: Alamos, 16 Sep. 1890, E. Palmer 665 (holo- type. US not seen: isotypes. GH-2 not seen. NY). Figures 10. 13a-f. Plants woody, with blackish brown bark, sparsely covered ascending; shoots perennial, basally along a single line with flexuous trichomes, 350— 450 um long; internodes 2.5-6 cm long, | mm i with 2 diam. Leaves with petioles 3-5 mm long, colleters at blade base: blades 25-55 X 7-15 mm. ovate, basally obtuse, apically acute to acuminate. adaxially sparsely covered only on veins and mar- 300—400 um long, covered, only on the midrib 300—400 um long. /nflo- rescences always one per node, 2- to 12-flowered, 2 gins with flexuous trichomes, abaxially isolatedly with flexuous trichomes, to 6 flowers open synchronously, sciadioidal (some- times a little elongated); peduncles 1—4 mm long. densely covered along a single line with flexuous trichomes, 300—400 um long: floral bracts 0.5—0.7 0.2-0.3 mm at the base, ovate to triangular, gla- brous; pedicels 2-5 mm long, glabrous. Floral buds 1.6-1.8 mm long. 0.8-1.1 mm diam., ovoid; calyx basally fused, campanulate, abaxially glabrous; ca- lyx lobes 0.8-0.9 x 0.4-0.5 acute; corolla campanulate, 1.7-2 mm long, abax- mm, ovate, apically ially white, glabrous, adaxially white, with verru- cose trichomes 100—120 um long on the apical and on the lateral parts of the lobes, and with down- ward-directed smooth trichomes 150-200 рт long on the central parts of the lobes: lobes basally fused, 0.4—0.6 mm wide, cally acute; gynostegial corona 0.5-0.6 mm long, erect to patent, ovate, api- shorter than to equaling the gynostegium; lobes erect, laminar, oblong. Gynostegium 0.1-0.9 mm high. 0.9-1 mm diam.. subsessile; anthers about as high as broad, rectangular, abaxially planar: anther wings 350—400 um long, extending along the entire length of the anther, anther wings of adjacent an- thers parallel to each other: connective appendages са. 250 jum long, ca. 350 wm wide, ovate, equaling the stamen in width, slightly to strongly inflexed: pollinarium with corpusculum ca. 120 jun long, be- tween 1.5 times and twice as long as broad, ovoid: caudicles 50-60 um long, cylindrical, geniculate, thickened at the insertion of the pollinium: pollinia subapically attached to the caudicles, ca. 140 jum long, 50-60 um wide, ovate in cross section, ob- 0.3—0.4 mm high, upper part 0-0.2 mm high, shorter than to longoid; style-head ca. 0.7 mm diam., equaling the lower part, flat to umbonate. Follicles and seeds unknown. Phenology. Collected in flower in March and April and from August to October. Distribution. Mexico: Chihuahua, Sonora; open places in mixed deciduous hardwood forests and tropical deciduous forests; 250-1200 m. Literature and illustrations. Wiggins (1940: 82, pl. 18). Metastelma latifolium and М. minutifolium con- stitute a pair of closely related, very small-flowered a corolla less than 2 mm long) species with different ecological preferences. Metastelma latifolium. char- acterized by ovate, apiculate leaves, shoots with one line of trichomes, pedunculate inflorescences, ovoid to oblongoid pollinia and flat to umbonate style-heads, is an inland and upland species, while M. minutiflorum, characterized by linear to slen- derly lanceolate leaves, shoots with two lines of tri- chomes, subsessile inflorescences, reniform pollin- ia, and conical style-heads, prefers habitats close to the coast. A detailed analyis of the tiny pollinaria and the style-heads is necessary for secure deter- mination (cf. Fig. 13). Additional м examined. MEXICO. Chihua- hua: Guasaremos, Rio Mayo, 10 Aug. 1936 (fl), Gentry 2373 (ARIZ). Sonora ca. 40 mi. NE й 1980 (fl). Lehto 24882 (ARIZ): Cer- ros Colorados: 34 a E of El Tabelo on rd. Verdes, 29 Oct. ‚ Van Devender et al. 95-118 (ARIZ); E Tesopaco, along new rd., р Аир. 1976 8, о 76-322 (ARIZ); Sierra de Ala- of Alamos, 31 Aug. 1996 (tl). Steinmann PU Sierra d Alamos, Arroyo Agua Escondida, 20 Aug. 1994 (fI), Fishbein et al. 1903 (ARIZ): Sierra de Alamos, Canon Chapotes 19 Mcr 1994 (fl), Fish- bein et al. 1606 ARIZ); along Hwy. 16, 3.5 mi. of epoca, 11 mi. E uf Puerto de la Sam 6 Sep. 1995 (fl). ИА 2460 (ARIZ): Rancho las Uvalamas, 25 Aug. 0 (fl), Martin & McWorther s.n. (NY); 26.9 km SE of is Yaqui on MEX 16 (Rancho 3 Mula), 21 Sep. 1997 (fl), Reina et al. 97-1087 (ARIZ, LL). 7. Metastelma das RE: O. Williams) ‚ Novon 7: 42. 1997. Cynan- О. Williams, Fieldi- . TYPE: pas: steep slope x Dus. us along Mexican Hwy. 190, 3 mi. S of La Trinitaria. alt. 5100 ft., 14 Oct. 1965, D. E. Breedlove & P. H. Ra- ven 13199 (holotype, Fl: isotype, DS not seen). Figures 14a-f. 15 iede & Mev chum conan e 19 ana, Bot. 32: 37. Mexico. Chia- Plants ascending: shoots perennial, basally 58 Annals of the Missouri Botanical Garden Yt ru 1005 D Figure 13. Metastelma latifolium and Metastelma и a-f: Metastelma sie hic a: Palmer 665, b-f: Steinman 955. a'-f': Metastelma minutiflorum, Gentry 4857. —а. Sho oot with inflorescence. —b, b'. Flow Corolla lobe, adaxial view. —d. d', Gynostegium and corona, —e, e”. Pollinarium. f f'. Style-head. Drawn by J. Conrad. Volume 91, Number 1 2004 Liede & Meve 59 Revision of Metastelma att EE (nr = ӨРЕ 1—7 — Zum AVI T is 2 а E vs 7 " rj qu "T TM on , ДУ Metastelma 2 oronatum. . Pollinarium. —f. Figure 14. d. Gynostegium and corona TEM re IOS d DN ^ c ا‎ 1745 к к view. . Corolla lobe. a. Shoot with inflorescences. —b. Flower. — Stvle-head. Breedlove & Raven 13199. Drawn by J. Com 60 Annals of the Missouri Botanical Garden So — [1 | ¡PES А E | | | | | P el^, * | o 4 ( | | —— == E 7 | L— XA T ~a | € AS | | lh { PN К | 1—4 | | \ ! | | | E A 31 WT SS Y ۱ NE AS. ال‎ 42 — — dy Е 30° OOCORWA PF e у 1 | لے‎ © | | к E \ ( ^h ө | | a x KR «LA 4. | yde di з г ее € | N | ai Ow › | * — — 4 — UA «а: = حا‎ — 0 „ | 25 NC м \ | l 2 | | | Nas c Ф | | Y Pit N AY e < | QUT RN e Е EN | | Е AY | E А A P [45 (^ А | Е س‎ а 7 20° B Metastelma longicoronatum کی‎ з A К. WE A | J | ) 2 ; A wt QUA SK M | Aj e Metastelma palmeri ар , „ 4 Matactal, f ; | n | S e e 1 y! * ( 0 Metastelma turneri x M. pen NS eo — Sa 5 | ao | „F EPIRI — = | < BUS: 15? -120* -115* -110° -105° -100° -95° -90° Figure 15. Distribution of Metastelma longicoronatum, M. palmeri, and M. turneri. woody, with dark brownish bark, densely covered smooth, downward-directed trichomes, 150-200 along a single line with flexuous trichomes, 250— 300 um long; internodes 2—6 cm long, 1 mm diam. Leaves with petioles 3-5 mm long, with 2 colleters at blade base; blades 20-40 X 6-12 mm, ovate, basally rounded, apically acute to acuminate, adax- ially isolatedly covered mainly on veins and mar- gins with appressed trichomes, 250—300 um long, abaxially glabrous. Inflorescences always one per node, 4- to 8-flowered, 3 to 5 flowers open syn- chronously, sciadioidal; peduncles 3-15 mm long, densely covered along a single line with flexuous trichomes, 150—200 jum long; floral bracts 0.4—0.5 0.2—0.3 mm at the base, ovate, with trichomes on the surface; pedicels 2-3 mm long, densely cov- ered along a single line with recurved trichomes, 150-200 um long. Floral buds 2—2.2 mm long, 1— 1.2 mm diam., ovoid; calyx basally fused, campan- ulate, abaxially with trichomes (mainly on the tube); calyx lobes 0.8-1 X 0.4—0.5 mm, ovate, api- cally acute; corolla campanulate, 2.5-3 mm long, abaxially white, glabrous, adaxially white, with ver- rucose trichomes, 100—150 um long, on the apical and on the lateral parts of the lobes and with шї long on the central parts of the lobes; lobes basally fused to ca. % of total corolla length, ca. 1 mm wide, patent, ovate, apically acute, recurved; gynostegial corona 2 mm long, longer than the gy- nostegium; lobes erect, apically slightly connivent, attached at the base of the stipe, laminar, elongat- edly triangular and with a conspicuously winged base. Gynostegium ca. 0.8 mm high, ca. 0.8 mm diam., on a stipe ca. 0.6 mm long; anthers about as high as broad, rectangular, abaxially planar; anther wings 0.6 um long, extending along the entire ength of the anther, anther wings of adjacent an- thers parallel to each other, in the same plane as the anther; connective appendages ca. 400 рт long, ca. 300 um wide, ovate, narrower than the stamen, slightly inflexed; pollinarium with corpus- culum ca. 160 um long, more than twice as long as broad, elliptic; caudicles ca. 70 jum long, flat- tened, slightly s-shaped, concave-convex; pollinia apically attached to the caudicles, ca. 180 jum long, ca. 80 um wide, ovoid; style-head white, ca. 0.5 mm diam., ca. 0.4 mm high, upper part ca. 0.15 Volume 91, Number 1 004 Liede & Meve 61 Revision of Metastelma mm high, shorter than the lower part. depressed- conical. Follicles and seeds unknown. Phenology. | Collected in flower in July. August, and October. Distribution. Mexico: Chiapas: cliff faces and limestone bluffs with seasonal evergreen forest: 1300-1700 m. Metastelma longicoronatum is a rare species en- demic to Chiapas. The elongated triangular shape of the corona lobes with a conspicuously winged, almost hastate base, is unmistakable. Additional specimens examined. A Sumidera, 22 km ^ of | . Breedlove 27183 (TEX); Comitán de Dominguez, 21 үч 1989 (fl). ад & Conrad 2557 (UBT). MEXICO. € T 19 Tuxtla Gutiérrez, 19 Aug. 8. Metastelma mexicanum (Brandegee) Fish- bein & R. A. Levin, Madroño 44: 270. 1997. Basionym: Melinia mexicana Brandegee, Zoé 216. 1905. degee) Bartlett, Proc. 632. 1909. TYPE: E of Culiacan, Basistelma mexicanum (Bran- Acad. Sei. 44: 30 mi. near Durango border, Cerro 3 Nov. 1904. J. S. Brandegee s.n (holotype, GH not seen). Amer. Mexico. Sinaloa: ca Colorado. Figures З. 1ба—К. . 1964. Bas- Report on 159 59. Cynanchum wigginsti Shinners, Sida 1: 36: ionym: — ja Torr. the U.S. & Mexica non Metastelma angosta Turcz.. kovsk. Obsc. Isp. Otd. Biol. 25: 314. 1 Melinia —1 (Torn) A. Gray, Proc. Amer. E Se i. 12: 73 dyra ê i (Tor Watson 1 15 Amer. Acad. Sci 60. 1380. Basis oom re An ; Marte гос. Acad. Arts 44: € : Mexico. Sonora: Santa Cris; Tow = ane 2x holotype, NY isotypes. GH!. MO not фм 1 Boundary Survey 2: 159. 1 Aca cl. cal Plants ascending. to 300 em high: subterranean organs constituting a woody rootstock: shoots pe- rennial. basally corky, densely covered along a sin- 200-250 um long; internodes 2-3.5 cm long. 0.5— gle line with recurved trichomes. | mm diam. Leaves with petioles 1-2 mm long, with 0 to 2 col- leters at blade base: blades 20-40 X 1-2 mm, lin- ear, basally cuneate, apically acute, marginally rev- olute when dry. adaxially isolatedly to sparsely covered on the midrib and the margins with re- curved trichomes, 200—250 рли long, abaxially gla- brous. Inflorescences always one per node. 2- to A-flowered, all flowers open synchronously, scia- dioidal; peduncles 0-2 mm long. densely covered along a single line with recurved trichomes, 150- 200 um long: floral bracts 2-3 * 0.2-0.3 mm at with trichomes the base. very elongate triangular. on the surface: pedicels 2-3 mm long. densely cov- ered along a single line with recurved trichomes, 150-200 um long. Floral buds ca. 3 mm long. ca 1.5 mm diam., ovoid; calyx basally fused: lobes 2.2-3 angular, calvx 0.3—0.4 mm, very elongatedly tri- apically acute, abaxially with recurved tri- chomes: corolla campanulate, 2.5-3.5 mm long. abaxially white, glabrous, adaxially white, adaxially with verrucose trichomes to 50 jum long on the api- cal parts of the lobes and with smooth trichomes, 150 pm long, forming a tuft at the entrance of the tube; lobes basally fused, ca. 1 mm wide, patent. ovate, apically acute, recurved; gynostegial corona ca. 1.2 mm long, shorter than the gynostegium: lobes erect, attached at the base of the stipe. at- tached with a distinct hook-shaped base, laminar, obovate. Gynostegium 1.4-1.6 mm high, 1—1.2 mm diam.. on a stipe 0.4 mm long: anthers broader than high. deltoid, abaxially planar; anther wings 300— 100 um long. extending along the entire length of the anther, anther wings of adjacent anthers par- allel to each other, in the same plane as the anther: connective appendages 900-1000 um long, 400— 500 pm wide, ovate, narrower than the stamen. erect; pollinarium with corpusculum ca. 300 рт long. more than twice as long as broad. elliptic: caudicles ca. 150 jum long, flattened, convexly re- curved, horizontal. ellipsoid-rectangular: pollinia subapically attached to the caudicles, ca. 300 jum long. ca. 150 um wide, ovate in cross section, ob- longoid; style-head white. ca. 0.8 mm diam.. ca. 1.2 mm high, upper part ca. 0.8 mm high. longer than the lower part, elongated-conical to obinfundibuli- Follicles 50-60 mm long, 4—5 mm diam.. obtusely deltate in form. one per flower. erect. fusiform, cross section, apically strongly beaked, medium light brown, seta side tuberculate, papillose with regu- brown, longitudinally grooved: seeds ovale. larly arranged papillae, aseta side sculptured with longitudinal ridges, with regularly arranged. papil- lae, marginally wingless, denticulate: coma 10-15 mm long. Phenology. Collected in flower from June to August and in October; in fruit in October. Distribution. Mexico: Sonora; U.S.A.: Arizona, New Mexico; open rocky slopes in eni wood- lands, tropical deciduous forest: 900-1550 m. Metastelma mexicanum is probably the most dis- tinctive Metastelma species in the southwestern U.S.A. and Central America. It possesses extremely large sepals reaching the length of the corolla ii — some specimens (e. g., Jenkins 93-88, ARIZ). The corona is basally fused to the stipe but also to the corolla base. a character shared only with a few South American Metastelma species (e.g.. M. par- 62 Annals of the Missouri Botanical Garden Figure 16. Metastelma mexicanum. —a. Habit. —b. Inflorescence. —c. Flower. —d. Sinus of corolla lobes, abaxial view, е. Two corolla lobes and one corona lobe, adaxial view. —f. Gy 5 E Pollinarium. —h. Style-head. ‚ Follic le. —j. Seed. a-h: Spellenberg & Repass 5353. i-k: Van Dann et al. s.n. Drawn by J. Conrad Volume 91, Number 1 2004 Liede & Meve 63 Revision of Metastelma viflorum, see introduction). Most diagnostic. how- ever, are the large. erect anther appendages. which are nearly as long as the elongated stvle-head. For a discussion of infraspecific variation, see Fishbein and Levin (1997) canum is probably М. arizonicum, judging from gy- The closest relative of M. mexi- nostegium structure. = Е \dditional specimens examined. MEXICO. Sonora: Sierra 5 оле road from Saguaribo to El Chiribo 23 Aug. 1993 ( Fishbein et al. 1362 ( а Т Mesa E ssopaco, above La Vinateria. ca. 35 К Alamos, 1275 m. 24 Aug. 1993, Fishbein el al. j ARIZ): ridge SW of Santa Barbara, 20 air mi. ENE 1450 m. 3 June 1993, ogi = 88 (ARIZ): Sierra | 148 of \lamos. Sierra de Alamos. ca. 900 m. 19 Aug. Steinmann s.n. (ARIZ): Palm Canyon, 17.7 mi.“ = и е in Cerro € inta de Plat side 1976. Van е et al. s.n. (ARI ca. | 200 m. ban n & $ (ARIZ). e D 1971. Van alios Goldberg & "Turner s.n. (ARIZ). U.S.A. New Mexico: Hidalgo Co.. Guade- lupe a. 16 Aug. 1979, Spellenberg & Repass 5355 Y 9. Metastelma minutiflorum Wiggins. Contr. Dudley Herb. 3: 71. 1940. elec: sono- rense Moran. in Levin & Moran. Vasc. Fl. Isla Socorro, Mexico (Mem. San Diego Soc. Nat. Hist. 16): 26. 1989, non C. minutiflorum N. Schum.. Nat. Pflanzenfam. 4(2): 1895. TYPE: Mexico. Sonora: hills between Magda- lena and Santa Ana. 7 mi. S of Magdalena. 14 Sep. 1934. J. L. Wiggins 7191 (holotype, DS not seen: isotype, USD. Figures 10. 134—£, Plants ascending. to 150 em high: shoots basally woody, with brownish bark. densely covered along two lines with recurved trichomes, 150-250 um long: internodes 2.5-5 em long. 1-1.5 mm diam. Leaves with petioles 5-6 mm long. with 1 or 2 col- (10—)20—50 × 3—5 mm. acute. leters at blade base: blades narrowly ovate or narrowly obovate. basally apically acute, adaxially sparsely covered on the midrib and the margins with flexuous trichomes, 250—300 um long, abaxially glabrous. /nflorescenc- es always one per node, 3- to 6-flowered. all flowers open synchronously, sciadioidal, subsessile: floral bracts 0.4—0.5 X 0.2-0.3 mm at the base. gular to ovate: pedicels 1—1.2 mm long. isolatedly trian- covered over the entire surface with appressed tri- chomes, 140-200 um long. Floral buds ca. 1.5 mm long. ca. 1 mm diam., ovoid: calyx basally fused, campanulate, abaxially glandular. ot with sparse trichomes: calyx lobes 1— ).3-0.4 mm. ovale, apically acute: corolla ыен 7-2 mm long. abaxially cream, glabrous, adaxi- ally 150— long. on the apical and on the lateral parts of the white. with smooth trichomes, 250 um lobes: lobes basally fused, 0.5-0.6 mm wide. patent to incurved, ovate, apically acute: gynostegial co- гопа 0.6—0.7 mm long, shorter than to equaling the eynostegium: lobes erect, attached directly under- neath the anthers, laminar, lanceolate. Gynostegium 0.7-0.9 mm high. anthers about as high as broad. 0.7-0.9 mm diam.. subsessile: rectangular, abax- ially planar: anther wings 0.4-0.5 um long. ex- tending along the entire length of the anther, anther wings of adjacent anthers parallel to each other. in the same plane as the anther: connective append- ages 300-350 um long. 200-250 um wide. obcor- date. equaling the stamen in width, slightly in- flexed: pollinarium with corpusculum. 75-80 pm long: caudicles 50 jum long. cylindrical. s-shaped. convex-concave, thickened at the insertion of the pollinium: pollinia apically attached to the caudi- cles. 75 um long. 50 um wide, ovate in cross sec- tion. reniform: style-head white, 0.3-0.4 mm diam.. 0.1—0.5 mm high. upper part 1.5-1.7 mm high. shorter to equaling the lower part. conical to knob- by. Follicles one per flower. 60-75 mm long. 4-5 mm diam.. fusiform, terete, apically strongly beaked. blackish brown. glabrous: seeds 5-6 mm long. 3-3.5 mm wide, pyriform. seta and aseta side tuberculate. papillose with regularly arranged. pa- pillae. marginally wingless, denticulate: coma 15- 20 mm long. Phenology. Collected in flower in April. Sep- tember, and November: in fruit in November. Distribution. thornserub and grassland: Baja California, Sonora: 200—500 m. Wiggins (1940: 82. Mexico: Literature and illustrations. 17 pl. ij. Cynanchum minutiflorum is a name not available for this taxon. because it is preceded by Cynan- chum minutiflorum K. Schum.. Nat. 4(2): 252. 1895, today considered a synonym of C. schistoglossum Schltr. (Liede. 1996b). Under Me- Metastelma minutiflorum is Pflanzenfam. tastelma, however, available and thus valid. For comments on a close relationship, see M. latifolium. Additional specimens e pow MEXICO. Baja Cal- ifornia Sur: canyon of W slope of peak ove кеч salt- Apr. 1962 (f. Moran 9141 Nov. (f), Gentry 4857 (NY, TEX): Yecora, 1998 n Tr Devender et al. 98-1285 (ARIZ). 10. Metastelma palmeri S. Watson. Proc. Amer. Acad. Arts 18: 115. 1883. Cynanchum gravi Shinners, Field & Lab. 20: 110. 1952. maccartil, Cynan- chum maccartii Shinners var. Sida : 360. 1964. nom. nov. for Cynanchum pal- meri (S. Watson) Shinners. Field & Lab. 19: 64 Annals of the Missouri Botanical Garden „% UY AA NOM ip ПОЛ 0.5mm а m ey чаш palmeri. 1. Shoot e inflorescence. —b. Inflorescence. —c. Flower. —d. Corolla кё. adaxial у ynostegium and corona. —f. Pollinarium. —g. Style-head. a: E 828. b. Pringle 6706. Liede & ыш 2612. Drawn by J. Conrad. 65. 1951, non Cynanchum palmeri (S. Watson) Metastelma macropodum Greenm., € Amer. Acad. Arts 181. 1898. Syr YP Blake, Contr. Gray Herb. 52: 83. 1951. TYPE: 33: 481. 1898. Syn. nov. TYPE: Mexico. Oaxaca: . . E . | Tomellín Canyon, 1540 m, 17 s 1897, C. G. Prin- U.S.A. Texas: Webb Co., Laredo on the Rio gle 7606 (holotype, NY!: isotypes, HBG!, M!, MO Grande, Aug. 1879, E. Palmer 824 (lectotype. not seen, Pl. SN. designated by Henrickson (1987: 98), GH not seen; isotype, NY!). Figure 17a-g. Plants ascending, 50-200(—400) cm high: sub- Volume 91, Number 1 2004 Liede & Meve 65 Revision of Metastelma terranean organs constituting a woody rootstock: shoots perennial, basally slightly woody. with vel- lowish brown to blackish bark. glabrous, isolatedly sparsely covered along a single line with re- 200—300 um long: mm diam. curved to flexuous trichomes, internodes 2—4 cm long. ca. | Leaves with petioles 2-7 mm long. with 2 colleters at blade blades (12>)16-35(255) X 2.5 ovate, basally rounded, apically acute, marginally base: —((-12) mm. revolute (when dry), adaxially isolatedly to sparsely covered, mainly or only on the midvein with ap- pressed to recurved trichomes, 150-250 pm long. abaxially glabrous. Inflorescences ен опе рег 3- to 8-flowered, 2 open synchronously, thyrsoidal to setadioidal: pe- node, extra-axillary, 2 to 6 flowers duncles 0-10 mm long. densely covered along a single line with recurved to flexuous trichomes. 150-200 um long: floral bracts 0.4—0.7 * 0.2-0.5 mm at the base, triangular, glabrous: pedicels 0—5 mm long, glabrous or isolatedly to densely covered 200— 250 um long. Floral buds 1.5-2 mm long. 0.8-1.5 along a single line with recurved trichomes, mm diam., ovoid: calyx basally fused. campanulate. abaxially glabrous or with a few sparse trichomes lobes 0.5-1.3 X 0.2-0.5 mm. ovate. apically acute to obtuse: corolla campanulate. 2.2— on the tube: J 3 mm long, abaxially white to cream, glabrous. adaxially white to cream, with smooth trichomes. 100(—150) рт long, on the apical and on the lateral parts of the lobes, apical trichomes not longer than lateral ones. and deflexed hispid trichomes on the central parts of the lobes: lobes basally fused, 0.0— | mm wide, patent, ovate, apically acute, recurved: gynostegial corona 0.7—1 mm long. equaling to lon- ger than the gynostegium:; lobes erect to inflexed. attached directly underneath the anthers, laminar. ovale to triangular. Gynostegium 0.4—0.7 mm high. 0.5-0.8 mm diam.. on a stipe 0.3-0.4 mm long: anthers about as high as broad, trapezoidal, abax- ially planar: anther wings 200-250 pm long. ex- tending along the entire length of the anther, the stamen forming a basal arch: anther wings of ad- jacent anthers parallel to each other, centrifugal. basally forming a distinct “mouth”: connective ap- pendages 200-250 um long. 250—300 um wide. ovale. narrower than the stamen. slightly inflexed: pollinarium with corpusculum. 100—150 um long. between 1.5 times and twice as long as broad, ovoid to elliptic: caudicles 50-70 um long. cylindrical. shaped. convex-concave: pollinia apically at- tached to the eaudicles. 120—150 рт long. ca. 50 um wide, ovate in cross section, oblongoid: style- —0.3 mm high. upper part 0.05—0.1 mm high, shorter than the low- head white. 0.4-0.5 mm diam.. 0.2 part, depressed-conical to umbonate. Follicles one per flower, 40-60 mm long, 4.5-5.5 mm diam.. i dark brown, glabrous; seeds 6—7 mm long, 3—4 mm wide, fusiform, terete, apically strongly beaked. ovate, medium brown, seta and aseta side smooth. marginally with 0.1—0.2 mm wide wing with entire margin. Phenology. Collected in flower from July to Oc- tober; in fruit in August. Distribution. Guatemala: Izabal: Mexico: Coa- huila, Nuevo León, Oaxaca, Puebla, Tamaulipas, Yu- catán; U.S.A.: Texas: on limestone, but also on ig- neous rock, matorral and thorn forest; 500-2000 т. Literature and illustrations. Henrickson (1987: 98, fig. Apparently, the same author published two re- placement names for Cynanchum palmeri, Cynan- chum grayi (Shinners, 1952) and Cynanchum mac- cartii 1964). the intentions of Shinners (1952, 1964) in both cas- (Shinners, As there is no doubt about es. and Cynanchum grayi would have been a name available for the taxon, an error is the most prob- able cause for this duplication. Henrickson (1987) remedied the situation by designating the same lec- totvpe for M. palmeri and С. maccartit. 9.336 9319 from the Yucatán Peninsula. both out of the Contreras from Guatemala and Conrad main distribution area in Mexico, display a corona a little more stout than usual, but are best accom- modated here until more material from this area vecomes available. The name applied to collections from central Mexico. specimens of Metastelma macropoda, . are indistinguishable from those of M. palmert. and M. M. macropoda is thus placed in synonymy of palmert. The flowers in the Metastelma palmeri complex are easily confused with those of M. minutiflorum and M. latiflorum. which show a very similar struc- M. latifolium and M. minutiflorum occur along the Pacific slope, while M. palmeri oc- ture. However. curs along the Gulf Coast. Morphologically, M. pal- meri can be distinguished by slightly larger flowers (corolla 2-3 mm long on average in contrast to less than 2 mm in M. latifolium and M. minutiflorum). broader and longer corona lobes (basally as broad as the anthers, and as long as or slightly longer than the gynostegium in contrast to smaller than the an- thers throughout and never exceeding the. gynos- tegium in M. latifolium and M. minutiflorum). and a more clearly stipitate gvnostegium. Most conspic- uous. though only visible under a dissecting micro- scope, are the very tiny pollinarta (120-150 X ca. 50 pm) with their thin and fragile caudicles. GUATEMALA. Iza- pr dditional specimens examined. Annals of the Missouri Botanical Garden bal: Cadenas (Puerto Mendez), 13 Oct. 1969 (fl), Con- treras 9336 (LL, NY). MEXICO. hog ge Canyon la yavia, in N side of Sierra de la Gavia, 2 Aug. 1973 (fl, fr), Henrickson 11724 (TEX): Catacol Mis. Aug. 1880 (fl), Palmer 828 (NY); 22 km ESE of La Cuesta del Plomo on the 2 اا‎ Boquillas Hwy., 7 June 1972 (fl), Chiang et 0 (NY, TE ~ al. 755 TEX): € че nda Mariposa, at the foot of the реА rn slope of the та de Puerto Santa Ana, 23 June 1936 (fl), Wynd & Ы pee (ARIZ, NY, WIS); Múz- Men Palm Canyon, 7 аси 1936 (tl). Marsh 305 (TEX): Rancho Las Tres Sabinas ca. 15 m. de Músquiz, 5 June 72 (f), Chiang et al. 7527A (T ; Rincón de Maria, 24 Aug. 1975 (fl), Wendt et al. 1328 155 ; Hwy. 57, 70 mi. N of Sails, 3 June 19 66 (fl), Wilson 11. 388 (TEX). Nuevo León: Mts. Bustamante, Aug. 1938 (fl), LeSueur 361 (TEX); Old Pasa road, Paso de O 8 pan 1989 (fl), Liede & Conrad 2612 (UBT). Oaxaca: Concepción qe sta, base del Cerro Pluma, Santa Lu- cia, 4 July 1994 (fl), Panero & Calzada 4026 (LL); distr. Huajuapan, 15 km al o de 55 13 July 1986 (fl), Mendoza & Mérida 2566 (LL); eforma entre Hua juapan de León y Meere 6 pe Pr 5 (fl). ou & Garcia 6656 (LL). Puebla: a Los Gri inea ca. 20 km al O de n 24 Sep. I 990 m тте 5790 (NY): ear Oaxaca, cerro de Costapa near San Luis Tultitlanapa, June 1908 (ib. Purpus 3244 (NY). agire 4.4 mi. NE of turnoff to биши, оп a ias (f), Nesom et al. 5957 (LL); Carretera eh vs 8 mi. 5 of "alm. 20 July ans (0), КО? TEX); along Hwy. 85, . М over rive brider at Colonia 1 1976 (fl). Stevens et al. 2260 (NY); 37 mi. out of Re eynosa on the 15 to San Fernando, 19 Oct. 1959 (fl), Johnston 4369 (TEX); near microwave tower above Villagran, 30 ded vlr (fb. : Ste- . Park, са 5 km N Celesttin, 16 May 1993, Conrad s (U BT), A. Texas: Brewster Co., end of trail down from Mt. 1 155 to camp in Basin, 2 Sep. 1947 (fl), ae 4088 (NY); Je ous Davis Co., Point of Rock on Ft. Davis— Valentine Hwy., 27 July 1947 (fl), Hinckley Pu A Live Oak Co.. Toad to Fiesta Marina just E of Rte. 534, 1.6 mi. N of Rte. 359, 26 July 1981 (f). Hill Da 7 (NY) 11. Metastelma pedunculare Decne., in A. P. de Prodr. 8: 1844. Cynanchum w woodsonianum L. О. Williams, Fieldiana, Bot. : 12. 1968, non Cynanchum pedunculare T 1786. TYPE: Guatemala. Cuesta de Piñ- ula, 1837, K. T. Hartweg 601 (holotype, G not seen; isotypes, NY—2!, P!). Figures 12, 18a-f. Candolle, ки vse ola Pittier, Contr. U.S. Natl. Herb. 13: 98. 910. Cynane ir ۰ ittier) L. O. Williams, En Bot. . TYPE: C osta Rica. San José: July pe " indue s.n. (holotype, US!; iso- type, P!) Plants ascending: shoots perennial, woody, with grayish bark, glabrous. Leaves with pet- ioles 4—7 mm long, with 0 to 2 colleters at blade base; blades 25-60 x basally rounded, apically acute to acuminate, adaxially gla- basally 7-17 mm, ovate, brous or sparsely covered only on the midvein with recurved trichomes, 300-400 um long. abaxially floral bracts 0.3-0.4 X glabrous. Inflorescences always one per node, 6- to 8-flowered, one specimen more than 15- to 20-flow- ered (Molina 23080), 1 to 3 flowers open synchro- nously, bostrychoid: peduncles 10-25 mm long, densely covered along a single line with recurved trichomes, 100—150 um long; rachis 0-2 mm long, straight or 3—17 mm long, angular (Molina 23080); са. 0.2 mm at the base, triangular, ciliate; pedicels 3—5 mm long. glabrous. Floral buds 1.5-2.5 1-1.5 mm diam., ovoid: calyx basally fused, campanulate, abaxially glabrous, ciliate or with very few trichomes; calyx lobes ca. 0.6 X ca. 0.3 mm, ovate, apically acute; campanulate, 2-3.5 mm long, abaxially cream, glabrous, adaxially creamish white, with verrucose trichomes, 100—150 um long. on the lat- eral parts of the lobes; lobes fused for ca. % of total mm wide, horizontal to re- mm long, corolla corolla length, са. 0.7 curved, ovate, apically acute; gynostegial corona 0.5-0.6 mm long, equaling the gynostegium; lobes inflexed, attached directly underneath the anthers, laminar, oblong. Gynostegium 0.3-0.5 mm high, 0.6-1 mm diam., on a column 0.6-1 mm long or only 0.3 mm long (Molina 23080); ie broader than high. rectangular, abaxially planar; anther wings 200—250 um long, extending along the entire length of the anther, anther wings of adjacent an- thers parallel to each other, in the same plane as the anther; connective appendages ca. 250 um long. ca. 250 um wide, ovate, narrower than the stamen, strongly inflexed; pollinarium with corpus- culum ca. 200 jum long, more than twice as long as broad, elliptic; caudicles ca. 150 jum long, ey- lindrical, s-shaped, convex-concave; pollinia ca. 75 200 um long, ca. 75 um wide, ovate in cross sec- tion, clavate; style-head white, 0.6-1 mm diam., 0.2-0.3 mm high, upper part flat. Follicles and seeds not seen, follicles 45 mm long (fide Standley, 2 Phenology. Collected in flower in July, August, and November. Distribution. Costa Rica: Cartago, Guatemala: Guatemala; Nicaragua: Estelí; volcanic soil, in ravines, thickets, along borders of fields: 1000-1800 m. Literature and illustrations. fig. 5) Specimens from Guatemala and Costa Rica show no constant differing characters, as Standley (1938b) observed correctly. He (Standley, 1938b) placed Metastelma sepicola in synonymy of M. pe- San José; Pittier (1910: 97, dunculare. In Metastelma pedunculare an exceptionally long and slender column (to 1 mm) carries the short and Volume 91, Number 1 Liede & Meve 67 2004 Revision of Metastelma С 0 y 3 b, Figure 18. Metastelma pedunculare. —a, a’, a". Shoot with inflore scence. ==”. Inflorescence. с. e”. Flower. = d. Corolla lobes, s view. n X Gynostegium and corona. —e - Pollinarium. —f, f'. Style-he ad. a-f: Specimens from Guatemala. a: Hartw 8, 601. : Molina & Molina 24843. a. A: Specimens from Costa Rica. a^: Tonduz s.n. b'— P: Taylor & Taslor 11393. a", b”: ы imen from Nicaragua, Mattia 23080. Drawn by J. Conrad. 68 Annals of the Missouri Botanical Garden flat gynostegium. The short laminar corona lobes are inserted directly beneath the anthers and are curved along the back of the anthers in a hook-like manner. In M. stenomeres, the only other highly stipitate species with the corona lobes inserted di- rectly underneath the gynostegium, the corona lobes are much longer than the gynostegium, while they barely exceed it in М. pedunculare. Molina 23080 from Nicaragua, a specimen cited by Stevens (2001) as representative of Metastelma species "B," is easily identified by its bostrychoid inflorescences, which are, however, more richly branched than in other specimens. Also, compared to material from Guatemala and Costa Rica, the stipe and the corolla tube are quite short. However, gynostegium and corona structure agree well with M. pedunculare, so Molina 23080 is included here until more material becomes available from which it can be judged whether the characters described for Molina 23080 are typical for Nicaraguan ma- terial and can be used to circumscribe a subspecies of М. pedunculare. Additional specimens examined. COSTA RICA. Car- go: 3 km SE of 5 11 Aug. 1967 (fl), Taylor cae (NY), 4285 (NY). Pur xe о Guacimal valley, Oct. 1984 (fl), Haber 575 5 "Т. L, WIS). San José: about : km NW of San Juan de Tibas song the Rio Virillo, 1 Aug. =e (9), 1 М е 11393 (NY); entre Frailes y La 1, 26 Nov. » (fl), ins 2617 (NY ^ Acosta, En- Pg 97.18 (U a GUATEMA yala: 7 km E of Guatemala City, 24 "e 1970 (fl), o 4189 (NY); betw. km 18 to 22 on Hwy. from Guatemala City to An- i Molina & T na 24843 (NY). > NICAR stelí: don › Estanzuela Creek, 8 km W of Estelf, 4 Nov. 1968 (fl), Molina 23080 (NY). 12. Metastelma пш А. ‚ Ргос. Ашег. Acad. Arts 21: 396. 1886. "s ynanc iin Lak Sai (A. Gray) cd rA 12: 91. 1987. TY Mexico. Chihuahua: near Chihuahua, 23 May 1885, C. G. Pringle 62 (holotype, GH not seen; isotype, US!). Figures 19a—f, 20 d. — кше iue isi (Scheele) Shinners var. breviflo- 1 S. Sida 1: 360. 1964. TYPE: U.S.A. Tex- as: wster Co., Big Bend oe Park, Chisos 18., 1800 m, 19 July 1952. G. L. Webster 4340 (holotype, SMU not seen). 100—400 cm high; subterranean organs constituting a woody rootstock (Hinckley 2105, 3106); shoots perennial, basally slightly woody, with yellowish brown or grayish bark, Plants ascending to decumbent, rarely glabrous, normally sparsely covered along a single thin line with recurved tri- 150-250 um long: 1-4 cm long, | mm diam. (green shoots) or 2-3 mm diam. (woody parts). Leaves with petioles 2-5 mm long, chomes, internodes with 1 or 2 colleters at blade base; blades 10—20(— 30) X 2-10 mm, ovate to elliptic, basally rounded, apically acute, marginally revolute (when dried), adaxially glabrous (except for a few isolated tri- chomes on midvein and the margin). abaxially gla- brous. Inflorescences 2- to 6(to 12)-flowered, 2 to 6 flowers open synchronously, sciadioidal; peduncles 0—2 mm long, densely covered along a single thick line with recurved trichomes, 150-250 рт long: floral bracts 0.6-1.2 X 0.3-0.6 mm at the base, triangular, glabrous; pedicels 1.2-2.5(—4) mm long, glabrous, sparsely covered along a single line with recurved trichomes, ca. 200 jum long. Floral buds calyx basally fused, abaxially glabrous; calyx lobes 1—1.2 X 0.4—0.5 mm, ovate, apically acute; corolla cam- 2.3-3 mm long, ca. 1.5 mm diam., ovoid: panulate, 2.8—4 mm long, abaxially white or yellow to brown (when dry), glabrous, adaxially white, with slightly verrucose trichomes, 200—700 um long, on the lateral and apical parts of the lobes, apically very long and wavy, and with hispid retrorse hairs, 300—500 um long, on the central parts of the lobes; lobes fused for ca. % of total corolla length, 0.7— 1.2 mm wide, patent, oblong to lanceolate, apically thickened, acute, with straight thickened margins: gynostegial corona 1.3-1.8 mm long, twice as long as the gynostegium; lobes erect, attached at the base of the stipe, solid to filiform, subulate, ex- tended into a long, slender tip. often incurled or coiled. Gynostegium 0.5-0.7 mm high, ca. 0.7 mm diam., on a stipe 0.5-0.7 mm long; anthers broader than high. wings ca. 250 jum long, extending along the entire trapezoidal, abaxially planar; anther length of the anther, anther wings of adjacent an- thers parallel to each other, centrifugal; connective appendages 200-300 um long, 200-300 jum wide, ovate, narrower than the stamen, slightly inflexed; 180 um long, more than twice as long as broad, elliptic; caudicles ca. 50 um long, cylindrical, geniculate, thickened at the insertion of the pollinium; pollinia apically pollinarium with corpusculum ca. attached to the caudicles, ca. 200 jum long, ca. 8 um wide, ovate in cross section, ovoid; style-head white to cream, ca. 0.6 mm diam., ca. 0.5 mm high, upper part ca. 1.6 mm high, shorter than the lower part, depressed-conical. Follicles one per flower, 40-62 mm long, ca. 5 mm diam., obclavate to fu- siform, apically strongly beaked, dark brown, glabrous; seeds 5-6 mm long, 2.5-3 mm terete, wide, ovate, reddish light brown, seta and aseta side smooth, marginally with 3 mm wide wing with entire margin; coma 20-25 mm long. Phenology. Collected in flower from May to Oc- tober; in fruit in April, and from June to October. Volume 91, Number Liede & Meve 69 04 Revision of Metastelma , 7 SAAN Hub NAH SUN N M CN МММ SS SS SSS — SSA Se — ey — | 0.2 mm м = ` === SE — — aa — — —c. Corolla lobe. adaxial view. —d. Metastelma pringlei. —a. Shoot with inflorescences. —b. Flower. inarium. —f. Style-head. a-f: Pringle 62. Drawn by J. Conrad. Figure 19 g 1 'stegium and corona. —e. Poll Gyn 70 Annals of the Missouri Botanical Garden | | T ; | | E: N | — | | | (C) Ke Й m~- | D E AN | | | i | } X — — po H p 35° | | | | | [AMA T | ` \ hi D Р | | à | | | | | | А ) | 1 | к 1] | IN ) 1 | | A \ \ { Li Јљ Фа i k ! 15 Re 30° V A у |. 4 | v4 * N v. — 1 25° | ry ү | E. 20* e Metastelma pringlei 7 " A А Metastelma schaffneri m mo EI 4 Гы» o — Pe 15° - | -120° -115° -110° -105° -100° -95° -90° Figure 20. Distribution of Metastelma pringlei and M. Distribution. Mexico: Chihuahua, Coahuila, Durango, Nuevo León, San Luis Potosí, Tamaulipas, Zacatecas; U.S.A.: gypsum and lava, matorral, open pine forest, scrub, roadsides; 500-2300 m. Literature and illustrations. 82); Henrickson (1987: 93, fig. The impressive crown formed by the long (to 1.8 mm), erect and often incurled corona lobes makes Metastelma pringlei an easily recognized species. The corolla indumentum with two different types of the conical gynostegium and the caudicles, Texas; rocky slopes, limestone, Wiggins (1940: C-). hairs, which are apically inserted to the conical pollinia via a thickened, convex area, demonstrate its close relationship to M. barbigerum (cf. Fig. la, b). Shin- ners (1964) even described U.S. material of M. pringlei as a variety under М. barbigerum. How- ever, the two species are easily and consistently separable by the corona lobes ca. twice as long as the gynostegium in M. pringlei (less than 1.5 as long in M. barbigerum). Only one specimen has been found that is possibly a hybrid between the two species (Mexico. Coahuila: 13 mi. S of Sabinas, 23 Aug. 1938, Shreve 8417, ARIZ). It displays the times schaffneri. slender leaves of M. pringlei in combination with a corona a little longer, but not twice as as long as the gynostegium and the bearded corolla lobes typ- ical for M. barbigerum subsp. barbigerum. Additional specimens examined. MEXICO. Chihua- hua: 4 km SSW of Cerro del Gringo in the Sierra del Diabolo, 30 Aug. 1972, Chiang et al. 9015f (TEX); hills near Chihuahua, 5 Sep. 1886, Pringle 955 (NY); Mts. NW of Chihuahua, 1 Aug. 1936, te ur 1053 (TEX); Santa Eulalia Mts., 1885, Pringle 70 (N Deis Sierra de la Campana, Rancho La Campana, 5 n 1977. Socorro Gonzales 840 (TEX): S slope & top of Sierra del Roque, NNE of Че 19 June 1973, Chic pida el ER 11387 (TEX): c l (air) mi. NE of с SW f above ee el Recuerdo in Sie 1973. Henrickson 12980b (TEX). Coahuila: Arroyo Seco, en el ‘ho el Tunal entre las Sierras San José y El Tunal, E de Parras de la Fuente, 14 Aug. 1983 (fl), . Rodriguez & Carranza 1111 (ARIZ, LL); ca. 12 (air) E of Boquillas in Sierra del RUM 10.4 (rd.) mi. NW of Rancho El Jardin, 27 July 1973 (fl). Henrickson 11497 (LL); Cañon San Lorenzo, SE d Saltillo, 2 Aug. 1975 (fl). Engard & Gentry 686 (LL); Canyon de Tinaja Banca, cen- tral mass of Sierra de las Curces W of Santa Elena 14 Aug. jme ). Johnston & Muller 261 (TEX); ien 6.9 1. SW of Cuatro Ciénegas on Hwy. to San Pedro, 27 Sep. 1986 (H. fr), pene 20400 (LL); 8 km W Vy MM Volume 91, Number 1 2004 Liede & Meve TA Revision of Metastelma Ciénagas, slopes of Sierra de la he га, 5 Aug. 1973 (fl). Ги et al. 12077 (NY. TEX): 35 (air) mi. W of in pe Canyon del la Шаш 3 (fl). Henrickson & дйн, 1202 28 Cuatro Ciénegas, de la Madera, 6 Aug. (LL): ca. 43 (air) mi. D of Cuatro Ciénegas, 4 mi. SW of Hacie ada Zacatosa, 7 Aug. 1973 (fl). He aki 12070 (LE) 52 mi. WNW of Cuatro Ciénegas (12 mi. E of La Vibora). 20 Sep. 1972. Henrickson 7891 (LL): ca. 07 (air) mi. SW of Cuatro Ci iénegas, above the main spring. 1.5 Ў, 15 Aug. 1973 (fl. fr). Henrickson . NW of | as Delicias, valle "V of Sierra « Е "nrickson E = Ly N of Ejido Piedritas, 12 5 Sep. 1972 e Chiang. et al. 930604 (NY) J: rd. from Esmeralda NE to San / s (via Mans and Emple us | Sep. 1940 (fl), Vim 4 Muller 834 (LL); 4 km E of Fraile, 12 July a ea E ер 362 (ARIZ. NY): E of Hacienda p Б l4 Jur 936 (fl), Wynd & Mueller 26 (ARIZ, NY): 11 km "E ч is и June 1941 (fl). Stanford et » km NNE of Las Margaritas in a large canvon E face i "ierra de = Е 24 Sep. 1972 (D. Chiang et al. 9501 (LE): c La Morada Ranch, 2 m. . Hinton 23343 (TEX): W base of Picacho del ида Vaione па. 23 Aug. 1941 ; Pore ‘ion Y W de la sierra la bap а ca. 45 a Carito Giz ey s. 5 Apr. Bounty et al. 17 (LL): Puerto de 1. fn: fous & Van Devender 9239 (ARIZ); SW flank of Sierra del Carmen, 15 Sep. 1972 (1). Chiang е! A 9274 (LL): 1.5 mi. S of Rte. 40 on road ph canyon, нар TAA rd. and Rte. 5 Aug. (LL): vicinity of im ho El е № a Y deed & close to the Chi- 1130 m Los re : 28 July (fl), Vilaseñor et al. 1621 (LL); N de of iit de la Fra- gua, E of Estacion Belloc, 38 km W of € uatro Ciénegas, 19 Oct. 1985 0» La vin et al. s.n. (LL); < Antonio, сапу‹ Antonio de los 2 Sep: 1940 (M. Johnston & Muller 950 ba L): Sierra de Parras. ca. 18 km S de Parras de la Fuente. 14 Oct. 1983 (fl, fr). Rodriguez 1024 ( ARIZ): Sierra San Marcos. W of Revn- olds Mine He quate rs, и Colorado turnoff W of San 2 July 1974 (fl). Engard. & Getz azaro Pass, Hwy. 57 Cienagas, (ARIZ); lower canyon & Mim e ES rra San Marcos op- posite Laguna Grande, 14 Aug. 5 (1). Reeves & Pin- kava 13075 (NY): S ih of Sierra Lus y Organos, ca. 9.5 km E of Puerto del Gallo, 8 Aug. 1973 (tl). Johnston el al. 12125B at Estacion Mic roondas Las age pan A me 1973 (fl). Johnston et al. 11457A (TEX); 26 mi. 30. 26 July 1958 (fl). Correll & Toto 20219 (LL 26 (air) mi. SW of Torreon. 6.8 (rd.) mi. NW of Hw . 40 toward Presa Francisco Zarca К Im 6 (fl), Henrick son & Prigge 15352 (LL). Nue 'vo León: ca. 15 mi. Matehuela "a IE 57, 24 July 1977 (fl, fr), алм el MÍ ТЕХ); La Becerra, 5 Sep. 1989 ( ton el s Г (LL); El Barrial, 4 Sep. 17660 (11): La Soledad, 26 Aug. 1940 (fl, fr), Shreve & Tinkham 9694 (ARIZ); Hacie Ya Pablillo, Gal- 1 t Aug. 1936 (fl). Taylor 172 (AR LL). San Luis — а * — T Cand. Potosí: 6.5 (rd.) mi. S of Arista. in Chihuahuan Desert. 6 Sep. 1971 (fl. fr), Henrickson 6427 (TEX): Ejido Cerrito Blanco, 10 km al E de Matehuala, 6 Oct. 1980 (fl). 11 (WIS): Charcas, July-Aug. 1934 pil Lundell (WIS). Tamaulipas: ca. 46 km WNW of Jaumave: 6.5 rd. km W of Miquihuana. Oct. 1982 (fl). He nrickson 19136b (TEX): 5 km al N de Estanque de los V km al Lemus 5517 alles o 25 NW de se beeen ty rife 1986 (f. Hernandez 1917 (TEX). Zacatecas: 5 (rd.) mi. NNE of Rancho Hidalgo. ca. 27 mi. E ‘amacho, in Chihuahuan Desert Sep. 1971 (fl. tr ji nrickson 6366b (LL). U.S.A. акне: ‘reek Canyon, Old Wildon Ranch, Big Bend Nat. ? E Sep. 1966 a), Correll, 33825 (NY): W side of Old Blue, Glass Mts., 2 June 1941. Warnock 21851 (NY): Big Bend, ee Trail, 7 Mar. 1989 (fl. fr) Liede & ies m (MO); Chisos Mts Ише 8115 . Pres di Co.. ca. | mi. is 1941 ^ "aM 2105 (N of Bats 14 Aug. 1941 (fl. fr), County not known: Valley of the Rio Ана А ) 3106 (NY). below Don- ana. Emory 1057 (I 13. Metastelma schaffneri A. Gray. Proc. Amer. Acad. Arts 21: 396. 1886. TYPE: Mexico. San Luis Potosí: near San Luis Potosí. 1876. J. G. Schaffner 052 (holotype, GH!: (frag- ment), NY!). Figures 20, 21a-g. isotype M i Lge lg S. poy Pr roc. Amer, Acad. Arts 1890. Syn. ne vw Mexico. Jalisco: mL RN 5 Мах 1888. E . Pringle 1776 (lec- pe, designated here. NY 002790571: isotypes, К ч pach, neg. 051460. GH not seen. MI. NYL A URS Plants ascending, to 200 cm high: shoots peren- nial, glabrous, sparsely covered along a single line with flexuous trichomes; internodes 25-30 em long. Leaves with petioles 1-2 mm long, with 2 colleters at blade base; blades 15-25 X 1.5-12 to elliptic to ovate, basally rounded, apically acute mm, linear to acuminate, marginally revolute (when dry). adax- ially sparsely, abaxially isolatedly covered only on the iris chomes, 5 um long. Inflorescences 4- to 6- midrib and the margins with recurved flowered, 2 to 5 flowers open synchronously, scia- dioidal, йы; and bracts 0.6-1.2 X 0.3-0.4 glabrous or with tri- mm at the base. triangular, chomes on the surface; pedicels 1-3 mm long, gla- brous. Floral buds 2-2.5 mm long. 1-1.3 mm diam., ovoid; calyx basally fused. campanulate. abaxially glabrous or with trichomes, mainly on the tube: calyx lobes 0.8-1.2 X 0.3-0.5 mm, ovate t triangular, apically strongly acute: corolla campan- ulate, 2-3 mm long. abaxially white with purple. glabrous, adaxially white, smooth trichomes, 150— 180 um long. on the lateral and on the apical parts of the V^ of total corolla length, 0.7-0.8 mm wide, patent, ovate, apically acute; gynostegial corona 0.6-0.8 mm long. longer lobes; lobes fused to Ca. than the gynostegium: lobes erect, attached directly 72 Annals of the Missouri Botanical Garden 7 Uu 8 | ji iM 9 2 17 +> — — Ñ — 0.5mm Figure 21. Metastelma schaffneri. n W a. Habit. —b. Shoot with inflorescences. —c. Flower. —d. Corolla lobe, adaxial . —e. Gynostegium and corona. —f. Pollinarium. —g. Style-head. a-g. Pringle 1776. Drawn by J. Conrad. Volume 91, Number 1 2004 Liede & Meve Revision of Metastelma underneath the anthers, laminar, ovate. Gynoste- gium 0.5-0.6 mm high, 1-1.2 mm diam., on a col- umn 0.3-0.4 mm long; anthers broader than high. rectangular, abaxially planar; anther wings ca. 400 um long, anther wings of adjacent anthers parallel to each other, basally widened, in the same plane as the anther: connective appendages 250—300 jum long. 350—400 um wide, ovate, narrower than the stamen, slightly inflexed; pollinarium with corpus- culum 100—150 jum long, more than twice as long as broad: caudicles 70—100 um long, cylindrical, slightly s-shaped, convex-concave; pollinia apically attached to the caudicles, 150—200 jum long. 80— 100 um wide, ovate in cross section, reniform: style-head green, 0.8-0.9 mm diam., 0.4—0.5 mm high. upper part 0.1—0.2 mm high. shorter than the lower part, depressed-conical. Follicles ca. 55 mm long, ca. 5 mm diam., obclavate, terete, apically strongly beaked, medium brown, glabrous: seeds ca. 15 per follicle, 6-6.5 mm long, 3 mm wide. ovate. sela and aseta side smooth, marginally with ca. 0.5 mm wide wing with entire margin. Phenology. Collected in flower in June, and from August to March; in fruit March, August, and September. Distribution. Mexico: Baja California, Durango, Guerrero, Jalisco, Morelos, Nayarit, Oaxaca, Pueb- la, Sinaloa, Sonora; hillsides and ravines, savanna, low open deciduous forest; 500-2900 m. Lectotypification of Metastelma multiflorum is necessary because there is no clue in the proto- logue as to where a holotype specimen was depos- ited. Of the many isotype specimens distributed, the NY 00279057 specimen is selected because it is accompanied by a nice drawing. Of the small-flowered Metastelma species with dense indumentum on the adaxial side of the co- rolla lobes. M. schaffneri can be confused with M. Metastelma schaff- neri is a species of western central Mexico, while M. schlechtendalii is distributed along the southern Gulf Coast, and M. turneri in northeastern Mexico. schlechtendalii and M. turneri. Metastelma schlechtendalii is best recognized by its pollinia, which are ovoid and attached to slightly geniculate caudicles, while the other two species have pyriform-reniform pollinia and s-shaped cau- dicles. Metastelma schaffneri has commonly much more slender leaves (to 12 mm wide) than M. tur- neri (to 18 mm wide) and a more slender gynoste- gium on a longer stipe (0.3—0.4 mm in contrast to 0.1—0.2 mm). Wiggins 15628, the only specimen from Baja California, with Metastelma agrees quite well schaffneri with respect to leaf indumentum, subses- sile inflorescences, corona exceeding the shortly stipitate gynostegium, and geniculate caudicles. However, corolla indumentum is very short (less than 100 um), and the corona lobes attach along the stipe a little distant from the anthers. Additional specimens examined. ifornia: Punta San Lorenzo, EUN mi. E of La Paz. 26 Nov. 1959 (fl), li cil 15628 (A urango: Durango & vicinity. 1896 (fl), үө 159 (NY). Gu MEXICO. Baja Cal- rie p near 5 1889 (8), Pringle 2749 (HBG). Mo ЕРИ Ng r- . 1895 (fl), 5 6205 (НВС. № PU 5). Nayarit: АБ quacatlan, on а S of town, 26 SA p (fl), Liede & Conrad 2962 (UBT). Oaxaca: 9 km al NE de Cuicatlán. rumbo а Concepci ión Pápalo, 23 Aug. 1980 . Mendraño 1630 (WIS); Santa Inez Mountains, 15 Oct. Mes 5626 (ARIZ, LL, NY): above Oaxaca, 7 ). Pringle 4766 (НВС, M. NY). Rancho San qe ‚ 10 a 65 NW de Molcaxac. 15 Oct. 1994 (fl). Tenorio et p: ) (LL); vicinity of due Luis Tultitlanapa; near wa b 1908 (fl). Purpus 2617 : Tehuacan, lava bed behind El Riego, 4 Aug. 1989 (fl), Linde & edge 2604 (MO). San Luis Potosí: 1879, ДЙ 313 (Р). Sinaloa: Feb. 1940 (fl), Gentry 5479 (MO, Кү oe А 8 Dec. 1939 (fl). Capadero, Sierra Tacuichamona. 12 Fe 1940 (fl), Gentry 5562 (ARIZ, NY): near roadside at Las Animas beach near Topolobambo, 28 Jan. 1964 (m. Pes s.n. (LL). So- T еу nora: Сайоп de Bavispe, Aug. 1940 (fl. White 5239 (NY). State unknown: 1905, Purpus 1259 (NY) 14. Metastelma schlechtendalii Decne.. in A. Р. de Candolle. Prodr. 8: 513. 1844, non Metas- telma schlechtendalii Chapm., Fl. South. U.S. 366. 1860. nom. Шер. Cynanchum schlechten- dalii (Decne.) Standl. & Steyerm. „ Publ. Field Mus. Nat. Hist.. Bot. Ser. : 226. 1947. TYPE: Mexico. Veracruz: Hacienda de la La- guna, Aug. 1828, C. J. W. Schiede 159 (lec- totype, designated by Stevens (2000: 256). P!: isotypes, HAL not seen, MO not seen, NYL W not seen). 14а. Metastelma schlechtendalii schlechtendalii. Figures 22e. Decne. var. M puce и Pittier, Contr. U.S. Natl. Herb. 13: nom. illeg.. non УНЕ 2 die in Cen Symb. Anull. 1: 249. 1899. TYPE: Ven Оше ‘hé: San O т. А ы: T. Heyde & E. Lux 5060 n S not seen; isotypes, F о seen, M!, NY!. UPS). Metastelma parv ani 5с ehh. nnaea 6: 731. 1831. m. illeg.. non 1 ar R. Br. ex Schult., in Roemer & Schultes, Syst. Veg. 6: 120. 1820. TYPE: Mexico. Veracruz: Hacienda de la La- guna, Sep. 1828, Schiede 260 (holotype. НАШ; iso- type, HAL!). Plants ascending, 80-150 em high: shoots an- Annals of the Missouri Botanical Garden Figure 22. Metastelma schlechtendalii. enicola, e: M. schlechtendalii var. Gynoste spo and corona. J. Conrac ore view. —d, d" ЗІ. : Molina 15672. Drawn by schlechte ote — € a VUE ENS padal anm 7 А CS * DANN Jy 7 4) E, Tow : M. schlechtendalti var. trichophyllum. d M. йн dice var. ar- a’. Shoot with inflorescences. —b'. Flower. —c'. Corolla lobe, e”. la —{'. $ lb a NA Molina m d“: Gentry Volume 91, Number 1 2004 Liede & Meve Revision of Metastelma rB o SA м | | : | m xp M | $ | q 3 " Е \ € |M ni 0 2 \ f ff A Dy a М \ / 1b | EE «NS OE S Tr | as . 3 \ р аі i N Ph f. } А gore X Plus r (oun | \, i g e a ) ps Pà l A.N ө | Ы ES ^ u а En \ f. e e e _ — — y ADAN = 6 EM е, a 20° ZA | LN Я SS Y Г = е at | — Ф. | nm SI — < | | à ? Ps „^^“ = к e “a | — - oan 5 KR 15° } A. ^a + Кы 3 me р x = — — + о Уй ло ө M. schlechtendalii var. schlechtendalii | у 4 M. schlechtendalii var. arenicola kg А M.schlechtendalii var. trichophyllum | | — | - 5? -110* -105* -100* Distribution of Metastelma schlechtendaliü. -115° Figure 23. nual, glabrous, sparsely to densely covered along a single line with recurved to flexuous trichomes. 150—300 um long: internodes 2-5 cm long. 0.5-1 mm diam. Leaves with petioles 1-5(-10) mm long. with O to 2 colleters at blade base: blades 15-15 X 4-15 mm, elliptic to ovate, basally cuneate to rounded, apically acute to acuminate, adaxially gla- brous or isolatedly to sparsely covered only on veins and margins (except for Aguilar 124 with iso- lated trichomes on the lamina). with appressed tri- chomes, ca. 200 pm long. abaxially glabrous or on veins and mar- gins with appressed trichomes, 200-300 um long. Inflorescences 2- to 12-flowered, all flowers open synchronously, sciadioidal, bostrychoid: peduncles (2-)4-15 mm long, glabrous, isolatedly to sparsely very isolatedly covered mainly covered along a single line with recurved tri- chomes, 200—300 рт long. Flowers very sweetly fragrant; floral bracts 0.5—0.7 X 0.2-0.3 mm at the base, ovate, glabrous or ciliate; pedicels 1.5-2.5 mm long. glabrous, isolatedly to sparsely covered along a single line with recurved trichomes, 100—150 ¡um long. Floral buds ca. 2.5 mm long, ca. 1 mm diam. ovoid: calyx basally fused, campanulate. abaxially with a few sparse trichomes mainly on the tube: calyx lobes ca. 1 X ca. 0.4 mm, ovate, apically acute: corolla campanulate, 2-3 mm long. abaxially white. glabrous, adaxially white, with verrucose tri- chomes, 150-200 um long, on the apical and on the lateral parts of the lobes: lobes basally fused. 0.6—0.7 mm wide, erect to patent, ovate to lance- olate. apically acute: gynostegial corona 0.8—1 mm long. slightly longer than the gynostegium: lobes erect, attached along the stipe. laminar, triangular. Gynostegium ca. 0.5 mm high, ca. | mm diam.. on a stipe 0.6-0.8 mm long: anthers broader than high. rectangular. abaxially planar: anther wings 150— 200 um long. extending along the entire length of the anther, anther wings of adjacent anthers par- allel to each other, in the same plane as the anther: connective appendages ca. 200 um long. ca. 150 um wide, ovate, narrower than the stamen. strongly inflexed: pollinarium with corpusculum. 120-150 um long. between 1.5 times and twice as long as broad, ovoid; caudicles 100-130 um long. evlin- drical, slightly geniculate; pollinia apically at- Annals of the Missouri Botanical Garden tached to the caudicles, 150-180 um long, 80—100 um wide, ovate in cross section, clavate; style-head white, 0.5-0.6 mm diam., ca. 0.25 mm high, upper part 0—0.1 mm high, shorter than the lower part, flat to umbonate. Follicles one per flower, 35—40 mm long, 2-3 mm diam., fusiform, terete, apically strongly beaked, dark brown, glabrous; seeds ca. 5.5 mm long, ca. 3 mm wide, oblong, light brown, seta side tuberculate, aseta side smooth, marginally with 0.5 mm wide wing with irregularly denticulate mar- gin; coma ca. 20 mm long. Phenology. Collected in flower from July to April; in fruit in November. Distribution. Belize: Corazal, Orange Walk, To- ledo; Guatemala: Guatemala, Jutiapa, Quetzalten- ango, Quiché; Honduras: Colón, Ocotepeque; Mex- ico: Campeche, Quintana Roo, Veracruz, Yucatán; very wide ecological amplitude, sand dunes, salt marshes, roadside vegetation, thickets, disturbed forests; sea level to 2000 m. Literature and illustrations. Pitter (1910: fig. 4а, b). Williams (1968) included Metastelma decipiens Pittier in the synonymy of M. schlechtendalii var. schlechtendalii because they share narrowly oblong- lanceolate corolla lobes that are adaxially densely barbellate, a corona attached along the stipe and slightly exceeding the gynostegium, and lanceolate or ovate-lanceolate leaves with trichomes only on veins and margins Metastelma sc hes htendalii var. schlechtendalii is a frequent species especially in lowland areas around the Yucatán Peninsula and up to 2000 m in Guatemala and Honduras. In the Yucatán, it is often difficult to distinguish from short-stiped, only vaguely barbate specimens of M. barbigerum var. liesnerianum, but the transversely oblong outline of the gynostegium is diagnostic against the conical gynostegium of the M. barbigerum complex. How- ever, there seems to be a lot of hybridization and introgression between the two species complexes especially in the Yucatán, judging from the numer- ous specimens with somewhat intermediate char- acters. Metastelma schlechtendalii var. schlechtendalii is easily confused with M. schaffneri and M. turneri, but can be distinguished by its conspicuously pro- truding anther wings (short and almost vertical in the other two species), and its flat-topped gynoste- gium with strongly inflexed anther appendages (rounded to slightly conical gynostegium with slightly inflexed anther appendages in the other two species Metastelma | schlechtendalii var. schlechtendalii has often been confused with M. parviflorum R. Br. ex Schult., a widespread taxon in northern South America and the Caribbean, but it is clearly distin- guishable by its only basally fused corolla and the corona lobes attached along the stipe, and not, as in М. parviflorum, at the base of the stipe, fused to Schlechtendal (1831: 731) commited exactly this error in his brief enumera- tion. Two years later (Schlechtendal, 1833: 518), he indicated the specimen on which he based his con- clusion, Schiede 260, collected at the same locality as the lectotype of M. schlechtendalii, Schiede 159. Decaisne (1844) noted the error, and TROPICOS takes the same view. the corolla. However, T specimens examined. BELIZE. Corazal: ca. 4 km SE of Sarteneja, 18 Mar. 1987 (f ч Dile ho Brant 32667 L, . Orange Walk: Honey Camp, Oct. 1929 (fl), Lundell 651 (NY). Toledo: Monkey ho Swansea branch, 26 Nov. 1941 (fl), Gentle 3815 (LL, NY). GUATEMALA. Guatemala: along Hwy. 18 about 7.2 mi. E of San José Pifiula, 16 July 1971 (fl), Stevens 1250 (NY, TEX). Jutiapa: Д 178 de а Арг. 1894 (fl), Heyde & Lux 63: : La Libertad, 2 June 1933, Lundell Pp (W IS); eee Road. km 82, 21 Nov. 196 , Contreras 7237 (LL); Santa Ana, 20 km al E de See Elena, 13 Nov. 1965 (fl), Molina 15672 (NY); al- rededor de Libertad, 30 km de Santa Elena, 22 ds 1970 (fl), Тип Ortiz 604 (NY); Santa Elena, en el camino para Santa Marta, a km 3, lado N, 12 Dec. 1970 (fl), Tún 0 1494 (NY); San Miguel, Savanna San Miguel, 3 km Мо San Miguel, 8 Dec. 1967 (fl), Contreras 7270 (LL); na Nat. Park, Bajo de Santa Fé, salida de Arroyo Corriental, 17 Jan. 1964 (fl), Contreras 3866 (LL). 5 Palmar, 13 Oct. 1934 (fl), Skutch 1438 (NY). District not known: St. Tomas, 1841, Friedrichsthal s.n. (NY). HO DURAS. Colón: 1.8 mi. strip on N bank of Río Guai- moreto, NE of Trujillo on old rd. to Castilla, 11 Dec. 1980 (fl), Saunders 740 . Ocotepeque: El Cerro, vi- San Шиш; 30 ae 1968 (i). Molina 22442 (NY); 9 Lempa river, rd. to Esquipulas, 29 Aug. 1968 (fl), Molina 22438 (NY). MEXICO. Campeche: alrede- dores de las grutas de Xtac arie a 3 km al O de Bolonchen de Hejón, 29 Sep. 1985 (fl), Cabrera & De e а NY); Zona "iiu. ч de Campeche, 28 Oct. 7 (fl), Gutiérrez 5462 (TEX). Quintana Roo: en la a ión a Laguna Ocum, a E km al Sur de GF. Carrillo Puerto, 13 Nov. 1980 (fl), Cabrera & Cortez 261 NY); Punta Bete, donde Jan. 1980 (fl), Sousa et al. Я Alto Lucero, Boca / 5 о 1971 (fl), Dorantes 385 T a Hacie de la со Schiede 11 (HAL); Хега- ‚ Wawra 625 (NY); Zac ‘марап and vicinity, Nov. 1906 (A), 7 ?urpus por (NY). Yucatán: 30 mi. E of Mérida at Hoctun, 7 Mar. 1973 (fl), Butterwick 78 (TEX); Uayma, ca. 20 km NW von Valladolid, 21 Nov. 1981 (fl, fr). oon & Geiseler 7016 (B), Feb. 1916 (fl), Gaumer 23436 f Progreso, E of port, ч 1938 (fl), Lundell & may 8113 TEX); along dirt rd. to Chamul SW of village, 14 Oc 1989 (fl), Mogensen 1151 (NY). State un. Berlandier $ s.n. ( p — — pm E — ж, Volume 91, Number 1 Liede & Meve 77 Revision of Metastelma Decne. var. TYPE: Mexico. Sonora: Mesa Masiaca, just N of the microwave tower, 6.5 km (by air) WNW of San José de Masiaca, ca. 200 m, 22 Nov. 1993, y W. 1 93-358 (with C. D. Bertelsen, A. Meyer, K. J. Johnson, M. M. Steinmann, A R. van Bere & F Wiens) (holotype. ARIZ!). Figures 22d", 23. 14b. Metastelma schlechtendalii arenicola Liede & Meve, var. nov. arietas varietati schlechtendalii similis, sed dif- fert d sn entibus subsessilibus, bracteis sepalibusque lanceolatis, corona staminali gynostegium multo super- anle. Plants ascending: shoots perennial, basally woody, with gravish brown bark. sparsely covered along a single line or over the entire surface (Gentry 5431) with erect or recurved trichomes, 200—250 um long: internodes 2—4 em long, 0.5-1 mm diam. Leaves with petioles 2-5 mm long. with 0 to 2 col- leters at blade base: blades 5—25(—50) X 4-10 mm, ovate to linear, basally rounded to cuneate, apically acute to acuminate, adaxially isolatedly to sparsely covered only on the midrib and margins with re- curved trichomes. 150-200 um long, abaxially gla- brous. Inflorescences always one per node, extra- 2 to 7 subsessile, rarely pe- axillary, 3- to 8-flowered, flowers open synchronously, sciadioidal, dunculate with peduncles to 4 mm long, densely covered along a single line with recurved tri- chomes. 150-200 um long; floral bracts 0.7-1 X 0.2-0.3 mm at the base. lanceolate, glabrous; ped- icels 1.5-3 mm long, sparsely to densely covered along a single line with recurved trichomes, 100- 150 um long, rarely glabrous. Floral buds ca. 2.5 mm long, ca. 1.2 mm diam.; calyx basally fused, campanulate, abaxially with a few sparse trichomes, calyx lobes 0.8-1.3 X 0.30.5 lanceolate, apically acute; corolla campanu- mainly on the tube; abaxially white, glabrous, 100—150 wm long. on the apical and on the lateral parts of the lobes: lobes basally fused, 0.5-0.7 mm wide, erect to patent, ovate, apically acute: gynostegial 2.5-3 mm long. adaxially white, with smooth trichomes. corona 1.2-1.5 mm long, distinctly longer than the gynostegium; lobes erect to inflexed, attached along the stipe. laminar, triangular. Gynostegium 0.7— mm high, 0.3-0.4 mm diam., on a stipe 0.5-0.6 mm long: anthers broader than high, rectangular, abaxially planar; anther wings 200—250 jum long, extending along the entire length of the anther, an- ther wings of adjacent anthers parallel to each oth- er; connective appendages ca. 500 um long. ca. 300 um wide, ovate, narrower than the stamen, slightly to strongly inflexed: pollinarium with cor- pusculum ca. 150 jum long, between 1.5 times and twice as long as broad, ovoid; caudicles ca. 80 um long. cylindrical, slightly geniculate; pollinia api- cally attached to the caudicles, ca. ca. 80 um wide, ovate in cross section, ovoid: style- 0.4-0.5 mm high. upper part ca. 0.2 mm high. equaling the lower Um long. head white, ca. 0.7 mm diam. part, distinctly umbonate or flat (Gentry 5431). Fol- 35-45 mm long, 4—5 obclavate, terete, apically shortly beaked, licles one per flower, mm diam., dark brown to black, glabrous; seeds 5-6 mm long. 3—4 mm wide, ovate. medium brown, seta and aseta side tuberculate, papillose with regularly arranged papillae, marginally wingless, denticulate: coma 20-30 mm long, yellowish. Phenology. Collected in flower in January, and from September to November: in fruit in October. Distribution. | Mexico: Sinaloa, Sonora: littoral sand hills, basaltic slopes, thornscrub and thorn forest; 30—400 m. In addition to the diagnostic characters of the new variety (subsessile inflorescences, lanceolate bracts and calyx lobes, and a corona considerably exceed- ing the gynostegium), (70%) have a slightly elevated style-head. while the style-head is flat to slightly umbonate in the other the majority of specimens two varieties throughout. The variety is restricted to the western Mexican lowlands, separated by at least 1000 km from the other two varieties. Gentry 5431 does not fit the description of Me- tastelma schlechtendalii var. arenicola perfectly: its shoots are sparsely, but evenly, covered with tri- chomes (in all other specimens, the trichomes only occur along a single, though occasionally rather broad, line), floral bracts and sepals are compara- tively short, the flowers relatively large, and the style-head is flat. Nevertheless. gynostegium shape and the protruding anther wings make it a member of the M. schlechtendalii complex and it is best ac- commodated here. Paratypes. MEXICO. Sinaloa: Altata, 31 Jan. 1940 (fl), Gentry 5431 (ARIZ, | pues abampo. 10 Oct. 1962 (fl), Skorepa 173 (WIS): nr. Yacht Hotel, e 5 Oct. 1964 (fr). Turner & Hastings 64-128 (ARIAL ra: summit of Cerro Prieto, vic. of microwave station, 15 km E of Navojoa above rd. 1 5 Sep. 1989 (fl). Sanders 9265 (ARIZ): Mesa Masiaca y vic inity, at summit of M.M. microwave tower, 6.5 km (by E of San Jose de Masiaca, 23 Sep. 1994 (fl), Friedman 379- 94 (ARIZ). Sono- 14е. Metastelma schlechtendalii Decne. var. Williams) Basionym: trichophyllum (L. Liede & Meve, chum trichophyllum L. O. Williams, Fieldiana. Bot. 32: 41. 1968. Metastelma trichophyllum comb. et stat. nov. Cynan- 78 Annals of the Missouri Botanical Garden (L. О. Williams) W. D. Stevens, Phytologia 64: 335. 1988. TYPE: › Telica, camino de Montaña Uval, 20 1903. A. Molina R. 13344 (holotype, Fl; isotypes, EAP not seen, NY! TEX!) Figures 22a'-f', 23. Honduras. Olancho: Cerca Plants ascending, to 500 cm high; shoots peren- bark, densely covered over the entire surface or along a nial, basally woody, with grayish brown single line with erect trichomes, 300—400 um long; internodes 2-5 cm long, | mm diam. Leaves with petioles 2—5 mm long, with 2 colleters at blade base; blades 15-35 7-15 mm, ovate, basally rounded, apically acute to acuminate, adaxially and abaxially sparsely covered over the entire surface with erect trichomes, 250—300 um long. /nflores- cences always one per node, (4—)6- to 12-flowered, 4 to 8 flowers open synchronously, sciadioidal: pe- duncles 5-15 mm long, densely covered over the entire surface or along a single line with erect to recurved trichomes, 300-400 um long; floral bracts 0.35-0.4 х 0.2-0.3 mm at the base, ovate, ciliate: pedicels 3-5 mm long, sparsely to densely covered over the entire surface with erect trichomes, 300— 400 um long. Floral buds 2-2.3 mm long, 1-1.2 mm diam., ovoid; calyx basally fused, campanulate; calyx lobes ca. 0.8 X ca. 0.3 mm, ovate, apically acute: corolla cyathiform to campanulate, 2.5—3(— 4) mm long. abaxially cream to yellow. glabrous, to 200 um long, on the apical and on the lateral parts of adaxially white, with verrucose trichomes, the lobes: lobes basally fused, 0.5-0.7 mm wide, patent, ovate, apically acute; gynostegial corona 0.8-1 mm long, equaling to longer than the gynos- tegium; lobes erect, attached along the upper third of the stipe, lanceolate, extended into a filiform tip. Gynostegium ca. О.А mm high, 0.8—1 mm diam., on a stipe 0.6-1 mm long: anthers broader than high, rectangular, abaxially planar: anther wings 200— 300 um long, extending along the entire length of the anther, anther wings of adjacent anthers par- allel to each other, in the same plane as the anther; connective appendages ca. 250 um long. ca. 150 um wide, ovate, narrower than the stamen, strongly inflexed; pollinarium with corpusculum. 120-150 um long, between 1.5 times and twice as long as broad, ovoid; caudicles 100-120 um long, cylin- pollinia apically at- 180-200 um long, 80-100 um wide, ovate in cross section, clavate; style-head drical, slightly geniculate; tached to the caudicles, white, 0.5-0.6 mm diam., ca. 0.25 mm high, upper part flat. Follicles (fide protologue) 25-50 mm long, 4-5 mm diam., obclavate, terete, apically strongly beaked, wingless, with dense indumentum: seeds not seen. Phenology. Collected in flower from August to November. Distribution. El Salvador; Honduras: Morazán, Olancho; Mexico: Chiapas; matorral, secondary vegetation, tropical deciduous forest; (500—)800— 2100 m en schlechtendalii var. trichophyllum is easily separated from the typical variety by the dense, equally distributed indumentum on stems and leaves. In addition, inflorescences appear to bear more flowers in variety trichophyllum, a char- acter hard to judge from herbarium specimens, as they might not have been picked at the peak of flowering. The distribution area of variety tricho- phyllum is disjunct between Chiapas (Mexico) and Honduras-El Salvador, with a gap in Guatemala, where the typical variety colonizes even the higher elevations. In Honduras, the typical variety and va- riety trichophyllum both occur, but variety tricho- phyllum seems to prefer higher altitudes there. However, as the floral characters fully agree with those of the M. schlechtendalii complex and the dis- tribution areas of the typical variety and variety trichophyllum are continuous to somewhat overlap- ping in Honduras, varietal (rather than specific or subspecific) rank for the taxon is appropriate, even though it is one of the easiest Metastelma taxa to recognize in the field by virtue of the equally dis- tributed indumentum on the leaves. ано specime ns examined. EL SALVADOR. San Salvador, 1271 (NY). HONDURAS. Mor- azan: е ЕИ ELC par Cerro de Hule 20 km Calderón ' of ا‎ 27 Oct. › (fl), Molina 18455 (NY): Ojojona, 45 km S de Терис al 28 a 1985 (fh). Aspra 65 (В); near Rio Guac pug & Los Laureles, NW of Te- gucigalpa, 4 Nov. 1966 (fl). inei ub (NY). Olan- cho: Carretera у pu del Río Juticalpa, 6 km de Ju- ticalpa, 18 Nov. 1963, Molina 13226 (NY. TEX): Valle Le rp entre El Zarzal y empalme a Galeras, i oS ч 3 (fl) Molina 13370 (NY). MEXICO. Chia rd. 1967 (ch. Ton 3092 from kl Bosque to Simojovel, 10 Oct. TEX); near microwave station of La Mina, 12 km S of X 190 nr Rizo de Oro. 16 Oct. 1971 e» Breedlove & Thorne 20658 (TEX); a 22 km al N de Ocozoc Pepe cam- mo a 0 24 Oct. 1985 (fl), Martinez 1428 (LL); 10 m SW of Ocosingo along rd. to San С deny 23 Sep. 1972 (fl), hu; оде 27820 (NY, ТЕХ); Puebla Nueva, 21 Oct. 1987 (fl), Pérez 85 (NY, TEX): Colonia Choro, 15 85 1966 io Ton 1548 (Т EX); La Roblada, Ocozocoautla, 15 Nov. 1971 (f. MacDougal H116 (NY); Las Rosas on rd. to ee ‘nustiano Carranza Aug. 1965 (fl). Roc et al. 997 (paratype, NY, WIS); 15 km SW of Suchiapa along rd. to Villa Flores, 3 Oct. 1972 (fl), Bree diu 26258 (NY, TEX). 15. Metastelma stenomeres (Standl. erm.) W. D. Stevens, Phytologia 64: 335. — & Stey- 1988. Volume 91, Number 1 Liede & Meve Revision of Metastelma — — Id > ME OS RATA, pt Vere ds 55 . Ee hate RA = Figure 24. d. Gynostegium and corona. —e. Metastelma stenomeres. a. Shoot Vis е -head. a-f: Schipp 674 Pollinarium. —f. . Flowe orolla lobe. pq =f . Drawn b J. Conra adaxial view. Annals of the Missouri Botanical Garden stenomeres Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 224. 1947. TYPE: Belize. All Pines, 2 m, 5 Feb. 1930, W. A. Schipp 674 (holotype, F not seen: isotype, NY). Figures 12, 24a— Cynanchum Plants ascending, 50-150 cm high: shoots pe- rennial, glabrous; internodes 4—6 cm long, ca mm diam. Leaves with petioles 4-6 mm long, with 0 or l colleter at blade base; blades 25-50 x 5-9 mm, slenderly elliptic or lanceolate, basally acute to cuneate, apically acute to acuminate, adaxially and abaxially glabrous. /nflorescences occasionally two per node, extra-axillary, 8- to 12-flowered, all flowers open synchronously, sciadioidal, but some- times basally dichasial; peduncles 3—5 mm long, sparsely to densely covered along a single line with recurved trichomes, 100-200 um long. Flowers sweetly fragrant; floral bracts 0.4—0.6 X 0.2-0.3 mm at the base, ovate, glabrous; pedicels 3-5 mm long. glabrous. Floral buds 3.5—4 mm long, 1.2-1.5 mm diam., slender ovoid; calyx basally fused to fused for ca. % of its length. very sparsely covered with trichomes; calyx lobes 0.6—0.8 х 0.2-0.4 mm, ovate, apically acute to ob- tuse; corolla salverform, 4—4.5 mm long, abaxially white, glabrous, adaxially white, adaxially with ver- rucose trichomes, to 300 um long, on the lateral parts of the lobes; lobes fused for ca. Y of total corolla length, 0.8-1 slenderly ovate, apically obtuse: gynostegial corona than the gvnostegium; lobes campanulate, abaxially mm wide, erect to patent, 2 mm long. longer erect, attached. directly laminar to filiform, lanceolate, extended into a long, slender tip. Gynostegium ca. 1.3 mm high, 0.7 mm without appendage of style-head, ca. 0.8 mm diam., underneath the anthers, on a column ca. 1.3 mm long; anthers broader than high, rectangular, abaxially planar; anther wings 0.3 um long, extending along the entire length of the anther, in the same plane as the anther; con- nective appendages 150 um long, 300 um wide. ovate, narrower than the stamen, strongly inflexed: pollinarium with corpusculum ca. 150 um long, be- tween 1.5 times and twice as long as broad, ovoid; caudicles ca. 150 um long, cylindrical, geniculate: pollinia apically attached to the caudicles, ca. 200 um long. са. 100 um wide, elliptical in cross sec- ca. 0.4 mm diam., ca. 0.6 mm high, 0.4 mm high. equaling or longer than the lower part, obinfundi- buliform or conical. Follicles one per flower, ca. 5X tion, pyriform; style-head white, upper part ca. = mm long. 4 mm diam., obclavate, terete, apically strongly beaked, light gray-brown, wingless, gla- brous; seeds unknown. Phenology. Collected in flower in February and from August to October. Distribution. Belize; in open pine forests, ham- mock edges; lowlands. A well-distinguished species endemic to Belize, which is most easily recognized by its floriferous inflorescences, the slender outline of the flowers, the very long (to 2 mm), acute and membraneous corona lobes, the long (to 1.3 mm) and slender col- umn carrying corona and gynostegium, and the usu- ally pronounced central elongation (to 0.4 mm) of the style-head. Additional specimens examined. BELIZE. Broken 1 8 mi. NE of Boomtown, 17 Sep. 1936, O'Neill 8660 (NY. WIS); near Manatee, 15 ч 1940 (fl), Gentle pad (NY, TEX); Monkey River, nr. Jenkins Creek, 1942 (fl), Саш 4190 (NY, TEX): Mountain Pi e > Ride, San к July-Aug.. 1936 (Н), Lundell 6656 (NY, TEX, ; Mullins River rd., 15 Nov. 1954 750 Gentle 8458 TEN. Traci i des k, Sibun River, 10 Aug. 1935 (fl), IS). TC un 1755 (NY, 16. Metastelma turneri Liede & Meve, nom. et stat. nov. Basionym: Cynanchum | maccartii Shinners var. latifolium Turner ex Henrickson, Sida 12: 98. 1987, non Metastelma latifolium Hose, Contr. U.S. Natl. Herb. 1: 106. 1891 TYPE: Mexico. Nuevo León: Las Anacuas, Mpio. Linares, 640 m, 5 Aug. 1980, С. S. Hin- ton 17935 (holotype, TEX not seen). Figures 15, 25a-f. Plants ascending, 150-200 cm high: shoots pe- rennial, basally slightly woody, with greenish to yel- lowish brown bark, sparsely covered along a single line with recurved trichomes, 200—350 um long: internodes 2—5 cm long, ca. 1 mm diam. Leaves with petioles 2-10 mm long, with 1 or 2 colleters at blade base; blades 10—40(—50) х 5-18 mm, ovate to lanceolate, basally rounded, apically acu- minate, marginally revolute (when dry), adaxially isolatedly to sparsely covered, mainly or only on the midrib and margins with recurved trichomes, 150-250 um long, abaxially glabrous or with a few recurved trichomes on the midvein. /nflorescences always one per node, extra-axillary, 3- to 8-flow- ered, 2 or 3 flowers open synchronously, sciadioi- dal, subsessile; floral bracts 0.4—0.7 X 0.2-0.3 mm at the base, lanceolate, ciliate; pedicels 2-4 mm long, covered along a single line with recurved tri- chomes, 200-250 jum long. Floral buds ca. З mm long, ca. 1.5 mm diam., ovoid; calyx basally fused, campanulate, abaxially glabrous or slightly verru- cose, ciliate; calyx lobes 0.5-1 X 0.3-0.5 mm, ovate, apically rounded: corolla subcampanulate, 2.54.0 mm long, abaxially creamish white, gla- Volume 91, Number 1 2004 Liede & Meve 81 Revision of Metastelma , 025mm , ' 05mm Figure 25. Metastelma turneri. —a. Sho Gynostegium with corona. —e ‚ Pollinartuin: f. Style-he brous, adaxially creamish white, with smooth tri- chomes, 50-200 um long. on the apical and on the than lateral ones, and scattered deflexed hispid tri- chomes on the central parts of the lobes; lobes ba- | i ovate to lan- sally fused, 0.7-1 mm wide, patent. d ot with inflorescences. —b. Flower. —c. ad. a: Hinon 17776. b-f: 7 75 11841. ateral parts of the lobes, apical trichomes longer rolla lobe. adaxial view. —d. Drawn by U. Meve. ceolate, apically acute; gynostegial corona 0.7-1 mm high, equaling to longer than the gynostegium: lobes erect, apically slightly reflexed, attached di- rectly underneath the anthers, laminar, ovate. Gy- legume 0.6-1 mm high. 0.5-0.8 mm diam.. on a stipe 0.1-0.2 mm long: anthers longer than broad. 82 Annals of the Missouri Botanical Garden trapezoidal, abaxially rounded; anther wings 200— 250 um long, not extending along the entire length of the anther; anther wings of adjacent anthers par- allel to each other, slightly centrifugal; connective appendages 200—250 um long, 250—300 um wide, ovate, narrower than the stamen, slightly inflexed: pollinarium with corpusculum. 100—150. um long. between 1.5 times and twice as long as broad, el- прис; caudicles ca. 100 um long, cylindrical, strongly s-shaped, convex-concave; pollinia apical- ly attached to the caudicles, 130—220 jum long, ca. 100 um wide, ovate in cross section, pyriform: 0.2—0.3 mm high, upper part ca. 0.1 mm high, shorter than the lower part, flat to umbonate. Follicles 45-55 mm style-head white, 0.5-0.7 mm diam., long. 5-6 mm diam., terete, apically strongly beaked, dark brown, glabrous; seeds 6—7.5 mm long, 3—4 mm wide, ovate, dark brown, seta and aseta side smooth, marginally with 0.5-0.7 mm wide wing with slightly irregular margin; coma ca. 20 mm long. Phenology. Collected in flower in May, July. August, and October; in fruit in October. Distribution. Mexico: San Luis Potosí, Tamaulipas; on dry hillslopes in scrub; 400— nm. Nuevo León, Literature and illustrations. Henrickson (1987: 98, fig. 1) Henrickson (1987) considered Metastelma tur- nert a variety of M. palmert, with which it shares the general shape of the corona. However, not only are the leaves and the flowers of M. turneri usually larger than those of M. palmeri (leaves: 16-35 X 2.5-7 mm vs. 10-40(-50) X 5-18 mm, ine up to 4 mm vs. up to 3 mm, respectively), but also the indument of the adaxial side of the corolla lobes in M. turneri is more reminiscent of M. schlechtendalii than of M. palmeri with the apical trichomes con- spicuously longer than the lateral ones. A new name has to be chosen for this species because Metastelma latifolium Rose (1891) has pri- orit y. One specimen (Mexico. bise rd. to Bus- tamente N of La Presita & 2.7 mi. N of Hwy. 70, 21 May 1982 (fl), Dorr 2359, TEX) shows the co- rolla lobe indumentum of Metastelma turneri, but the leaf shape and flower size of M. palmeri and is assumed to be a hybrid between the two species. Additional specimens 5 MEXICO. Nuevo León: К! Chipinque. ca. 6 km al sur de ino а la е 7 Oct Monterrey, cam- . 1998 (fl. fr). lane al & Carranza s.n. (TEX); Linares, 12 May 1980 (fl), Rzedowski 17776 ТЕХ); ера jos А Monterrey, 17 Aug. 1903 (fl). Pringle 11841 (TEX); Montemorelos, 10 Aug. 1988 (f). Patterson 6476 m Villa Santiago, Cañon de Potrero ои A July — (fl). scd 2099 (NY, TEX). ye is Potosí: 75 km N of S uis Potosí, on side rd. t mal 'ázar at km 11 . 19 July 1982 (fl), Fryxell 3820 ). Tamaulipas: Tm km NW of Palmillas on P to ahs 1 8 14 Aug. 1941 (fl), Stanford et al. 920 (N 2341); Mpio. Victoria, Altas Cumbres, 29 June е i 3 1534 (TEX). EXCLUDED NAMES Metastelma angustifolium Turez., Bull. Soc. Imp. vaturalistes Moscou. 1852(2): 315. 1852. TYPE: Mexico. Miradores, alt. 3000 ped.", J. Linden 1353 (holotype, P!). A member of the genus Orthosia. Metastelma eulaxiflorum (Lundell) Liede, Novon 7: 41. 1997 Wrightia 5: ¿ynanc hum ا‎ Lundell, 351. Jobinia eulaxiflora (Lundell) W. D. e Novon: 244, 2000 TYPE: Guatemala. Baja Verapaz: Union Bar- rios, high forest hilltop, E of km 154, 8 June 1975, C. L. Lundell & E. Contreras 19401 (ho- lotype, LL!; isotypes, MO not seen, S! Transferred to the previously exclusively South American genus Jobinia E. Fourn. by Stevens (2000) because of its paired, axillary. diffusely pa- niculate inflorescences. Metastelma 5 Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 956. 1938. Cynan- chum pe (Standl.) L. O. Williams, Fieldiana, Bot. 32: 36. 1968. TYPE: Rica. Entre San Ramón y La Palma de San Ramón, June 1928, A. M. Brenes 6179 (holo- type, F not seen: isotype, NY!). Costa This species is the only presently known member of the genus Scyphostelma Baill. (formerly *Cynan- chum" sect. Microphyllum) in Central America be- cause of its basally fused corona and its character- istic branching pattern of long shoots and short shoots bearing distichously arranged leaves. Metastelma glaberrimum Woodson, Ann. Missouri Bot. 107: 17. 1937. Cynanchum glaber- rimum (Woodson) L. O. Williams, Bot. 32: 36. 1968. TYPE: valley of the upper id Chiriquí А пеаг Monte Lirio, ca. 1830 m, 11 July 19 Seibert 300 кен 0. bas. 5 по! seen). Gard. Fieldiana, Panama. Chiriquí: A member of the genus Orthosia because of its basally fused corona and adaxially glabrous corolla lobes. Volume 91, Number 1 2004 Liede & Meve Revision of Metastelma . Williams) W. D. Ste- 1988. C ynanchum miserum ^r О. Williams. Fieldiana. i 38. 1968. TYPE: Guatemala. Alta Verapaz: Im да in i Pansamala, alt. 1260 m. July 1887, Н. v. Tuerckheim 1290 (holotype, US!; isotypes. P^. Metastelma miserum (l. € vens, ra "M F not seen, GH not seen, A member of the genus Orthosta because of its basally fused corona and adaxially glabrous corolla lobes. Metastelma pubescens (Greenm.) W. D. Stevens. 405. 1983. Astephanus pubes- Acad. Arts 32: 209 Morelos: wet bar- above Cuernavaca, 2100 m. 21 Sep. C. C. Pringle 6507 (Fl, GH! P! MO not NYD: "in the mountains near Cuerna- ” 3 Aug. 1896, C. C. Pringle 7203 (GH!). Phytologia 53: cens Greenm.. Proc. Amer. 1897. SYNTYPES: Mexico. ranca 1896. seen, — vaca. A member of the genus Orthosia because of its adaxially papillate (but not. indumented) corolla lobes. green stems. double inflorescences, and twin follicles. Metastelma rubens (L. O. Williams) W. D. Stevens. Phytologia 64: 335. 1988. каш = rubens L. O. Williams. Fieldiana. Bot. . 1968. TYPE: Guatemala. a ia Santa Elena. 2400-2700 m, 27 July 1933, A. К Skutch 503 (holotype, US!; isotype, NY!). А member of the genus Orthosia because of its basally fused corona and adaxially glabrous corolla lobes. Metastelma sepium (Decne.) W. D. Stevens. Phyto- logia 604: 335. 1988. Vincetoxicum sepium Dec- ne.. in A. P. de Candolle. Prodr. 8: 526. 1844. Cynanchum tipi (Decne.) Standl.. Contr. U.S. Natl. Herb. 23: 1177. 1924. TY PE: Mex- ico. Oaxaca: in hac 2300 m, Н. G. Gal- eotti 1533 (holotype. Pl; isotypes, Gl, K!). A member of the genus Orthosia because of its basally fused corona and adaxially glabrous corolla lo yes. INSUFFICIENTLY KNOWN NAME Metastelma selerianum Schltr.. Bull. Herb. Boissier 6: 841. 1906. TYPE: Mexico. Chiapas: Cerro de Tonalá. "in pratis montanis." Feb., C. Seler & H. Seler 2056 (holotype. В destroyed). In contrast to Metastelma lanceolatum. no spec- imen annotated “Metastelma — selerianum™ by Schlechter or anyone else was found, and the de- scription gives no hint as to the identity of the tax- on. Literature Cited Albers. E & U \sclepiadoideae. and e -volutionary б е within Apocynaceae s.l. Missouri Bot. 24—050. mima & n Mev ve, 1990. IOPB е И some hii 2: Asclepiadaceae. ТОРВ Newslett. 15: 11— | 4. Bartlett. H. H. 1909, VII. Descriptions of Mexican phan- товат». Proc. Amer. Acad. Arts 44: 630-037. „ 1844. In: R. B. Hinds (editor), The Botany f the Voyage of H.M.S. ا‎ Smith. Elder, London. Blankinship. J. W. 1907. Plantae III. Annual Rep. Missouri Bot. can 123-223. Brown. R. 1810. On the Asclepiaeae, a Sale Order of Plants Separated. from. the кок of Jussieu. R. Brown, London. Chaturvedi, S. К. 1988. Abiotic M tion in p hirsuta Wight (Asc "reis skle epi 93-02. D A cy, W. G. 1987. Meve. 2001. A karvological survey of Periplocoideae and Sec E ае. =j = — . Par a of Panama—C hecklist ui чуя * Pari 1: “The Intro- u tion and Checklist. Monogr. Syst. Bot. Missouri Bol. LE: Decaisne M. J. à due ^ 1 p Vol. 8. Treuttel. Wü Fishbein, M. Martin, D. "i T. R. Van Devender & R. Wilson b mw E mir s Río Mayo a Univ. Tucso R. Levin. cle q ae): A new the status of Basistelma Bartlett. Fournier, E. 1885. Ascle TE eae. In: tius (editor), Flora > ie n ‘nsis, Vol. pographia Regia, Müncher Notes on Cynanchum (Asclepiada- О. ‚ Asclepiadaceae. Pp. 490-084 in A. Prodromus Systematis Naturalis P: ^ Arizona Press 1997. Me ie in mexicanum (As- combination and re-e ores "i е 14: 208-2 ^P von e ae 298. Ty- Henrickson, J. 1987. ceae). Sida 12: Hewson, Н. J. 1988. Plant indumentum—A hi mdbook of к, Austral. Fl. Fauna Ser. 2: 1-2 Kunze, H. 1991 poe EN and function in asc da ‘piad pol- E D Syst. Evol. 176: 227-253. 19 . Corona нер nectar system in Asclepiadi- nae (Asc ا ا‎ eae). Flora 192: 17: 3 8 Levin, G. X. & огап. 1989, Asc le din eae: Rubi- aceae, The Vascular Flora of Isla Socorro. Mexico. 16 ed.: 26; 50-51. Society of Natural History, San Diego. Liede, S. 1996a. Sarcostemma (Asclepiadaceae =A con- troversial generic. circumsc 5 reconsidered: Mor- P m л idence. Syst. Biol. 3144. — \ revision E Cma | cei РЕ! in и а. Ann. Missouri Bot. Gard. 83: — 1997. Subtribes and genera of the tribe CN (i чч ynaceae—Ascle spiadoida ae)—A synopsis. pu M 241. i rs. 1994. Tribal cr np of Asclepia- daceae ge nera. Taxon 43: 201—2: .unze. 2002. un and the Cynan- chinae (Apocynaceae—Asclepiadoideae)— A molecular. anatomical and latex triterpenoid study, Organisms. Di- versity Evol. 2: 239-209. A. Täuber. 2000. Ascle к Asc de. Ree Sarcostemma R. Br. (Apocy- 84 Annals of the Missouri Botanical Garden naceae—Asclepiadoideae)—A controversial generic cir- cumsc jose às с EET E : bae е from trnL-F spac- rs. P ). vol. 25: & 0 ircumscription of the genus Cynanchum (Арес упас раг ‘lepiadoideae). Syst. Bot 301. Lumer, C. . Yost. 1995. The reproductive biology of Vincetoxicum nigrum (L.) Moench (Asclepiadaceae), a Mediterranean weed in New York State. Bull. Torrey Pome Asclepiadaceae, Atlas of Medicinal и of Middle Хей ica: MN to Yucatan. Charles Thomas, Springfield, Ilinc BL Н. F. 19 ae—New or interesting le aM 13: 96-116 1. Asclepiadoideae (Apocynac све) da cad- hs Gerais, Brasil. Bol. Bot. Univ. Sao Paulo 19 169. ‚ М. W. Chase, b. J. Goyder & J. Griffiths. 2003. eae classification: Evaluating the phylogenet- World Asclepiadoideae (Apo- Адорна ic no of New cynaceae). Taxon 52: 33-50, Salvin, 0. 1869. A synopsis of p сеш Clothilda. Trans. Entomol. Soc. uer 675 397. Schlechtendal, D. E Hora Insulae Sti. India Occidentalis. iue : 722-7 = 33. De plantis mexicanis A. G. Schiede M. e. collec v шо adfert. Li o 8: 513-528. Se hlec he H. ). Ascle р ас pe i: 1. Urban ety Sy ger 1 Vol. * erlin. 1906. Asc ed Herb. Boissier 6: Thomae, "Bull. 839-844. Schumann, К. 1895. Asclepiadaceae. /n: A. Engler & К. Prantl lior p natürlichen Planzenfamilien. Vol. ‚ 1: 189-305. Engelmann, Leipzig. Shinners, L. H. 1952. нан grayi Shinners nom. nov. os E Lab. 20: . 1964. Texas Asc E СЯ eae other than Asclepias. Sida, р = 1: 358-367. Standley, P. J Apocynaceae; U.S. a Herb. 23: Asclepiadaceae. Contr. 1166-1194: 168( 30. 19 e Asclepiadaceae. Pp. 177-17 Standley Calderón (айй), Lista 1 de las e de El Salvador. Tip la Union Dutriz her- manos, pe Salvador. 8a. Asc Vide ee In: Flora of Honduras. Publ. Field ib Nat. Hist., Bot. Ser. 17: 387-388 . Ase Vial eae. /n: Flora of Costa Rica. Publ. re Mus. Nat. Hist., Bot. Ser. 18: 949—960. Record. 1935, Asclepiadaceae. In: The Мену ae flora of British 1 Publ. Field Mus. Nat. Hist., E )-331 Stevens, W. D. New names and saree in Apocyneae, * 1 ae. Phytologia 64: 333-335. ———. 2000. New and interesting E И (Apocy- naceae, Ascle HEN n Novon 10 v ds —— ———. 2001. Ascle e eae. In: W. ens, C. Ul- O. M. Montiel ics Flora de 0. Monogr. Syst. Bot. Mis- loa Ulloa, A. Pool & & Nic 'aragua, Tomo 1: 234-27 souri Bot. Gard. 85. Sundell, E. 1994. Asc Vʒ‚p аи family. J. Arizona-Nevada Acad. Sci. 27: 169-18 Thomas, A. V. & Y. Dave. 1991. 5 and phy- logenetic significance of the colleters in the family Apo- „ Feddes Rep. 102: 177-182. 1986. Part И (6): Collectors "S." In: F. R. Stafleu Бао). Index Herbariorum. The Hague. Weberling. F. 1989. Morphology of Flowers and Inflores- cences. ACE Univ. Press, Cambridge. ). New and 1 known species from the Sonoran 2 8 Contr. Dudley Herb. 3: 65-86. ———. 1980. Asclepiadaceae. Pp. 202- ae in Flora of Baja California Stanford Univ. Press, Stanforc Williams. L. . Asc M ri In: Tropical Amer- ican Plants X Fieldia ana, Bot. 32: 35-61. е Tropical American 5 9 XII. Fieldiana, 102. R. E. 1941. The North American Asclepiada- . Ann. Missouri Bot. Gard. 28: 193-244. ooa. APPENDIX I. Numerical index of species. la. М. arizonicum subsp. arizonicum lb. М. arizonicum subsp. chiapense 2a. M. barbigerum var. barbigerum 2b. M. barbigerum var. m 2c. M. barbigerum var. veracruzens За. M. californicum e ы ш ЗЬ. M. californicum subsp. lanceolatum 4. M. cuneatum 5. М. eliasianum 6. M. latifolium 7. . longicoronatum 8. M. mexicanum 9. M. minutiflorum 10. M. palmeri 11. V. pedunculare 12. M. pringlei 13. V. schaffner l4a. M. schlec endi var. schlechtendalii 14b. М. schlechtendalii var. arenicola 14c. M. schlechtendalii var. trichophyllum 15. M , . Stenomeres 16. М. turneri APPENDIX 2. Index to collections examined. The numbers in parenthe ses refer to the corresponding species in the text and in the Numerical List of Species presented above in ie dix 1. Aguilar 124 (14a); Albers Е al. 46448 (2а); Andréz кеже, & Carranza 1111 ; Aspra 65 (14c). Jarkelew 217 (3a); Bakley 13360 (2a): Barkley et al. 155 (2a); Barlow 28/17 (2b); Bennett et al. 646 (12); жш s.n. (За); Berlandier s.n. (14a); Blake 7793 (1b); Boege 862 (3b), 1301 (2c), 1438 (3b); Brandegee s.n. (3a), (4); Breedlove 14010 (2b), 27183 (7), 27820 (14c), 28258 (14c), 28986 (1b), 62261 (3a), 62528 (3a); Breedlove h Raven 13199 (7); Breedlove & Thorne 20658 (14c); 2984 (2c); Burgess 5695 (la): Burgess & Van Doce 5235 (12). 7661 (3a); Butterwick 78 (14a). ;abrera & Cortéz 281 (14a); Cabrera & De Cabrera 9505 (14a); Calderón 1271 (14c); Calvert 117 (2a); Car- ranza & García 1080 (12); о ѕ.п. (2с); Сһаѕе 7418 (3b); Chiang et al. 7527A (10), 7550 (10). 7964 (2а), 8161р (2a), 9015 (12), 9274 (12), 9366a (12), 9501 (12), 11387 (12); Conrad 9319 (10); Contreras 532 (2b), 3068 3866 (14a), 6602 (14a), 7237 (14a), 7270 (14a), 8466 (2b), 9336 (10); Correll 33825 (12); Correll & Cor- rell 12756 (2a); Correll & Johnston Wee да), 20219 (12); Cowan 3616 (2a); Cowan & Mag 0 (2c); Cow- an et al. 5: jos (2a); Croft 62 (2a); С! 75 1 1125 (2a), 2984 (2a), )ОО (2a). Tae hie 406 (2b); Davidse & Brant 32667 (14a); Volume 91, Number 1 2004 Liede & Meve 85 Revision of Metastelma Davis & ое 66297 (3a); Dorantes 5270 (2с); Dorr 2359 (1( 16); Drees et al. 74-15 (la): Duke 12414 (5) Dwyer 2800 (5). Edwards s.n. (2b); \ (11); Engard & Getz 331 (12): Engard & Gentry 686 (12): Ertter 4906 (2a). 5357 (2a). Felger 9579 (1a), 9635 (3a), 15588 (1a); Felger & к; үт 15620 (la) Felger & Quijada-Mascarenas 91-3: : Felger & Schne idis 95-84 (la): Felger & E A | (la), 84-490 (la); Fernández 4731 (ЗЬ): Fernandes & Barkley 14485 (2a); Ferris 8589 (3a); Fishbein 2400 (6). 2759 (4); Fishbein & McMahon 2750 (3a). 2709 (3a): SH ein et al. 1362 (8), 1448 (8), 1606 (6), 1903 (6). 1905 (6): Flügel & Geiseler 7016 (14a): Fried- man 015-95 (4), 104-95 (4). 150-95 (4). 199-95 (4). 2 94 . 319-94 (14b); Friedrichsthal s.n. (14a): сид ll 3651 (2, а). 3820 (16); Fryxell & Anderson 3617 (3b) Galeotti 1571 A (3b); Gaumer 539 (2b). 23436 (14a). 24230 (14а); ди 1508 (2b). 1755 (15). 2517 (Ab): Gentle, P. H. (15), 3815 (14a). 4190 (15). 8458 (15): Gentry 1098 TR 1298 (la). 2373 (6). 4857 (9). 5184 (13), 5431 (14b), 5479 (13), 5562 (13), 6723 (2a). 7136 (3a). 7438 (3a). 7716 (3a). 14304 (la), 14466 (la): Gentry & Engard 23247 (12); Ghiesbreght 664 (1b): Gilman | 10 (la): Ginzbarg 131 (10); borde 76-322 (6); Goodding 116-45 (1a); 208-45 (1а); Gould et al. 2804 man 386 (2b): Gregg s.n. (2a): ee 1632 (2b). 14a). Haber 615 (11); Hall 521 (2а); Harmon 4189 (11); Har- tweg 601 (11); Hastings & Turner 67-64 (3a); Haynes & Stewart s.n. (За); Heller 1559 (2a); Henrickson 6100 (12). 6366h (12). 6427 (12). 7891 (12). 11497 (12). 11724 (10). 12070 (12). 12481 (12). 12980b (12). 19138b (12). 19303 (2a), 20400 (12); Henrickson & Lee 17559 (2a): Henrick- & Prigge 15352 (12): Henrickson € Wendt 12028 : Hernandez 1534 (16): 1562 (2a). 1917 (12). 1940 : Heyde & Lux 3060 (14a). 0348 (14а): Hill 10627 (10): Hine kley 2105 (12), 3106 (12), 3954 (10), 4088 (10): Hinton 17660 (12), 17776 (16). 23343 (12): Hinton el E 1310 (3b), 19616 (12), 19732 (2a), 24186 (2a), 242 (2a): Howell cogo За). — ~ с = Se =. ш. — = =, la): Green- 5462 11): Johnston, 1. М. Jenkii -88 (8): Jimenez 26 17 ( 8438 (12) Nu I. M. & Muller 261 (12). 834 (12). 950 (12); Johnston. M. С. 4369 (10). 5063B (2a); John- ston, M. г & Graham 4525 (2a). 4674 (2a): Johnston. М. et al 157A (12). 11991 (10), 12077 (12). 12125B Keil 4323 (la): 2 & Smith 14531 (5). LaBounty et al.! 12); Lavin et al. s.n. (12); Le Roy s.n. (la): Lehto е P Lemus 11 (12); LeSueur 361 (10): 1053 (12): Liede 3242 (2b): Liede & Conrad. J. 2557 (7). 2004 (13). 2610 (2а). 2612 (10). 2613 (10), 2955 (За). 2961 (13), 2962 (13); Liede & Meve 2500 (2a). 2501 (12): Lindheimer E а, 992 (2a) Lundell 651 (14a). 3502 (14a). 5517 (12). 6050 (15). 8734 (2a). 16709 (2b). 17022 205 L dun 11 : t m 118113 (14а). 1 116 (116): Machuca 7400 (3b): Marsh : Martin s.n. (За); Martin & Mc- Д sey 3034 (13). 14284 (146); Mar- 5 (2a): Matuda 3765 (2b); McGee .n. (12); О n & Bowers 2119 (3a): Modos & Mé ant 2566 (10); Mendraño 1630 (13); 51 (14a); Molina 3 (1b). 13162 (Ib). Pee (14c). 13344 (14c). 13370 (14). 155 (14c). 18607 (14c). 22438 (14а), a). rs (11): Molina & Molina 24843 (11): Moore Jr. & Wood Jr. 4409 (3b). 4259 (3b): Moran 9141 (9); Moser et al. 18152 (la) : Mueller, C. Н. 2099 (16), 8115 (12): 1 sr, C. H. & Mueller, M. T. 188 (16): Müll- er, F. 1853 a & d зе 10076 (la): Nesom et al. 5957 (10) Nixon & Cowan 831 (3a). O'Neill 8659 (2b). 8660 (s " Palmer 159 (13), 22: . 360 га, 626 (la). 665 (6). 824 (10). 828 (10); Panero & Calzada 4026 (10). 4040 (10): Patterson 647 6 (16), 6519 (2a): Pennell 10220 (2a). 10320 (2a), 17729 3539 (5): Pringle 70 (12), 9 2836 (2а), 3138 (3b). 4766 (13). 6646 (2a). 6706 (10). 10838 (3b). 1 yes (13), 11841 (16). s.n. (la): Purpus 1259 (13), 2094 (2c. 14a). 2617 (13). 3244 (10). 7273 (Ib). 9078 (Ib), 9167 (3b). 9169 (1b). 10217 (Ib). Reeves & Pinkava 13075 en Reina et al. 96-286 du in 1087 (6); Reverchon 1558 (2a); Rodríguez 1024 (12 {ое & Roe 1885 (3b); Roe et al. 997 (14c): Rose 1 : vr 1218 (12); = Il et al. 11402 (la). 11584 (la). . 12115 7592 (2a). = دن‎ = c — a) Ф Ф N ^^ ее oy А pend = 2 їл mm Э: — = ás 5 00 & Tenorio 3294 (3b): Sand- ers 9265 (14b); Sauer & ire 3137b (2b): = rs 740 (Ida): Schaffner 652 (13); Schiede 11 (14a), 159 (14a). 200 (14а); Schipp 674 (1 5) Shortman 90-27 s ‘Shre "se 6468 (3a). 7197 (3a), 8417 (2a X "y 312 (3b); Shreve & Tinkham 9694 (12): Skorepa 173 b Skutch 1438 (14а); Small & Wherry 11875 (2a): si M124 (2a); So- corro Gonzales 840 (12): Sousa et al. 10929 (14a); Stan- ford et al. 68 362 (12). 920 (NY id (16): Stein- mann 955 (6). 03-358 (14b). s.n. (6. 8 Varela 972 (la): Spellenberg & Re Des 5353 (8): Stevens 1250 (14a). 1: “es (2с); Stevens et al. 2259 (10). 2260 (10): Stewart 466 (12 Taylor, J. & Taylor С. 11393 (11), 12532 (2c); М. 172 (12); Taylor. R. J. 4251 (11). 4285 (11): Tenorio . 7660 (13), 10441 (3a); Tharp, B. C. 248 (2a), (2a). s.n. La): Thomas & Felger 11785 (1a). 11942 " a) Ton 1548 (14c). 3092 (14c): Tonduz s.n. (Herb. (11): Torres & Garcia 6656 (10); Tourney s.n. (Та); Тип m ot (14a). 1494 (14a); Turner, B. L. s.n. (13): buts H. & Hastings 64- E (14b): К. М. et al. 79- ve na Tyson et al. 2 (5). Van Devender & UE s.n. (8); Van Devender & Reina 98-1132 (la): Van Devender & Yetman 94-698 (1); Van Devender et al. 92-121 (3a), 92-133 (4), 93-1253A (4), 93-1265 (4), 93-1284 (4), 93-1310 (la). 93-65 (la). 98- 1285 (9), 95-1189 (6), 84-29 (la). s.n. (8): Vilasefior el al. 1621 (12): Villarreal & Carranza s.n. (16): et al. 3115 (2a). Wagner et al. 4120 (12): Warnock 21851 (12): Wawra 625 (1 aa) Webster & Webster 119 (2a); Webster & Wilbur ): Steinmann & Taylor, ‘Turner, Villarreal 3060 (2a): зу et Е 2 (10): White. D. & Mott 74 (3b): White. 5. 3239 (13): Wiggins 7191 (9), 7491 (la). 15048 (За), : 508 5 dm 15628 (13): Wilson 11388 (10): Woodson et al. 1263 (5). 1499 (5), 1500 (5): Wright 1676 (2a). 1077 e s.n. (2а); Wyatt 1464 (2a); Wynd & Mueller 26 (12), 244 S. coll. ым bl 3198 (2a). APPENDIX 3. Index to scientific names. Astephanus pubescens Greenm. Basiste Ima 1 (Jorr. ' an 1t 61 randegee) Bartlett € Cynanchum arizonicum (A. Gray) 8 39 barbigerum (Scheele) Shinners 43 86 Annals of the Missouri Botanical Garden barbigerum (Scheele) Shinners var. breviflorum Shin- barbigerum Scheele var. veracruzense Liede & Meve 46 californicum (Benth.) Moran 47 californicum Benth. chiapense (А. Gray) Standl. & Steverm. 42 californicum Benth. subsp. californicum 47 collinum (S. F. Blake) Standl. & eig 42 californicum Benth. subsp. lanceolatum | (Schltr.) Liede & Meve 49 5 A. Cray 42 num S. F. Blake 5 Brandegee 52 decipiens Pittier 73 ners y 3: filisepalum (Standl.) L. O. Williams 82 glaberrimum (Woodson) L. O. Williams 82 grayi Shinners 63 infimicola L. O. Williams 54 eliasianum Dugand 54 liesnerianum L. O. Williams 46 eulaxiflorum (1 un 1 iede 82 эш oronatum L. O. Williams 57 filisepalum Stam maccartii Shinners var. latifolium Turner ex Henrick- glaberrimum Age - son 80 infimicola (L. O. Wiliams W. D.Stevens 54 maccartii Shinners var. maccartii 63 lanceolatum Schltr. 4 miserum L. O. Williams 8: latifolium Rose 57 palmeri (S Satan) Blake 04 liesnerianum (L. O. Williams) Liede : longicoronatum de O. Lo lliams) L le « Meve 57 sc Fu ы (Decne.) Standl. & Steyerm. 73 sepicola (Pittier) L. O. W = 66 sepium (Decne.) Standl. ¿ miserum (L. O. Williams) W. D. Stevens 83 4 palmeri 5. Watson 63 sonorense Moran 63 xirviflorum R. Br. ex Е һин. 73 * . . he stenomeres о & Steyerm. 80 je ол. Schltdl. 7: tric ida di pcd dont L. О. | Williams 77 pedunculare Decne. m wigginsu Shinners 6 pringlei A.Gray 6 woodsonianum = O. Williams 66 pubescens ones ) W. D. Stevens 83 Ditassa mexicana Brandegee 49 rubens in 12 9 M. D. Stevens 83 Jobinia a undell) W. D. Stevens 82 scha 71 йт сег иЗ De спе Me linia angustifolia A. Gray 6 schlechtendalii Decne. var. arenicola Liede & mexicana Brandegee 61 Metastelma oo = 39 albiflorum S. Watson : Meve 77 schlechtendalii Decne. var. schlechtendalii 73 а Miel üm Tore E Ki eas | us n var. trichophyllum (L. ми Тигез. 82 ams) Liede бг Meve 74 selerianum Schltr. 83 arizo гау 39 i "OM Ру sepicola Pittier 66 arizonicum A Gray subsp. arizonicum 39 T E \ Gray subsp. ehiapense (A. Gray) Lied sepium (Decne.) W. D. Stevens 83 : Cas nba. е e (A. Cravi Liede І | ечеи т satay suas] I 7:8) “ee stenomeres (Standl. & Steyerm.) W. D. Stevens 80 & Meve 42 trichophyllum (L. О. UA W. D. Stevens 77 barbigerum Scheele 43 turneri Liede & Meve barbigerum Scheele var. barbigerum 43 atsonianum Standl. E barbigerum Scheele var. liesnerianum (L. O. Wil- Éatahas angustifolius (Torr. A: = Watson 61 liams) Liede & Meve 46 Vincetoxicum sepium Decne. 83 A TAXONOMIC REVISION OF Hong De-Yuan? and Pan Каг- Yu? THE РАЕОМА ANOMALA COMPLEX (PAEONIACEAE)! ABSTRACT Based on field observation, examination of over 350 she els of exsiccalae. and investigation of all the relevant types. a taxonomic. revision of the Paeonia anomala complex is presented. Typifications are clarified for P anomala. P. intermedia, P. hybrida, P. sinjiangensis. P. altaica, and P. veitchii. The identity of three well-known taxa. P anomala. P. hybrida. and P intermedia, is also clarified. Two species. P anomala and P intermedia, are recognized, with the circumscription of P anomala emended to include P veitchii as its subspecies. Paeonia sore includes two sub- species: subspecies anomala and subspecies veitchii with the former distributed in Central Asia, Siberia. and the northeastern I uropean part of Russia, while the latter occurs in China southeast of the Gobi rm Porania intermedia is distributed in Central Asia. but with an isolated locality in Georgia. Paeonia sinjiangensis is treated as a synonym L. The of P anomala, and P. hybrida Pallas is treated as a synonym of P. tenuifolia > lectotypes of Paeonia laciniata. Paconia beresowskii, and Paeonia intermedia are designated here in the present paper. Key words: Paeonia, Paeonia anomala complex. Paeonia hybrida. Paeonia intermedia, Paeoniaceae. The Paeonia anomala L. complex (Paeoniaceae) needed a comprehensive taxonomic revision and comprises a group of herbaceous peonies in Central review of the nomenclature. For this purpose, the Asia. Siberia. and adjacent northeastern European first author examined all available specimens of regions. They are characterized by leaves biternate this group in the herbaria at BM. HNWP. K, LE. with leaflets decurrent at the base, leaflets finely NWTC. PE. SHI. SHMU. TBI. XJBI. XJNU. and segmented. with segments of a lower (the best de- NJU (see Appendix 1). In addition, the types of PF veloped) leaf ranging from 70 to 100 in numberand anomala. P. hybrida, P. intermedia, and the other 4-32 mm in width, and by bristles along veins on six specific and varietal names were examined the upper blade surface. The only species with LINN (Herbarium of the Linnean Society). BM. К. which it may be confused is Paeonia tenuifolia |... and PE. respectively. but the latter has leaves even more finely seg- mented. with segments more than 130 in number HISTORICAL REVIEW and 0.5-6 mm in width. ps * pi eE Within the Paeonia anomala complex the type Since the late 1970s. two new species in the : : м ES › | species, P anomala, was described from Siberia by Paeonia anomala complex have been described д ih à i : : . ere i . пра , Linnaeus (1771). Pallas (1789) described three ad- from Xinjiang, the Central Asian part of China: P sinjtangensis К. Y. Pan (1979) and P. altaica K. M. Dai & T. H. Ying (1990). For a better understand- ing of the species group in Xinjiang, we made an ditional species, P. laciniata and P. sibirica from Siberia. and P hybrida from a plant raised from seeds of P. tenuifolia cultivated in the Botanic Gar- Su. x den of the St. Petersburg Academy in Russia. expedition in 1993 to the Altai and the Tienshan „ dat MR 8 е : Paeonia sibirica Pallas (1789) shares an illustration in Xinjiang and found that morphological features ars os (tabula) with Р, laciniata, but does not have a de- of the group in Xinjiang were not consistent. with the N by Schipezinsky (1921, 1937) and Gamaulova (1961); Stern’s (1946) treatments and nomenclature are different from theirs but do not scription. In his monograph of Paeonia, Anderson (1818) recognized only P. anomala, reducing P. hy- brida as a synonym of P. tenuifolia for the first time. De Candolle (1818) recognized P anomala. P. hy- reflect the reality of the group in Xinjiang, either. : x р de J brida, and P. laciniata, but later (1824) treated P Thus. we realized that the P anomala complex The authors are grateful to the National Geographic Society for financial support (Grant 6408-99), which allowed us to examine herbarium specimens at the Komarov Institute of Botany, the Russian Academy of Sciences. and to conduct fie е in Georgia. The project is also supported by the Nat ional Natural Science Foundation i i hina ( NSEC Grant 30130030). We are grateful to the curators of the following herbaria: BM, НМР, N. LE. N „EE, SEI. SHMU. i XJBI. XJNU. and: XJU. We sincerely thank Zhou iang for his assistance with fie VE pet li Ol 10- Ling and Ma Li-Ming for their help in preparation of the manus Laboratory of Systematic and Evolutionary Botany, Institute of M The Chinese Academy of Sciences, Niang- shan, Beijing 100093, China. hongdv(Ons.ibcas.ac.cn ANN. MISSOURI Bor. GARD. 91: 87-98. 2004. 88 Annals of the Missouri Botanical Garden laciniata as a synonym of P anomala. Meyer riety anomala and variety intermedia (Pan, 1979). (1830) described the fourth species, P. ae from the Altai Mountains. Ledebour (1842) cite anomala, P. hybrida, and P. intermedia, is 5 laciniata as a synonym of P. anomala. Trautvetter (1860) recognized only one species, reducing P. hy- brida as a variety of P. anomala, and treating Р. intermedia as a form of variety hybrida. Lynch (1890) cited P. hybrida and P. anomala, and stated that the former was native to the Caucasus. Huth ( omala, 892) adopted the widest species concept of P. an- which includes four varieties: var. typica, var. hybrida (= P. intermedia), var. nudicarpa, and var. emodi (— P. emodi, confined to the western Himalayas). A further treatment by Krylov (1901) divided the complex into two species, P. anomala and P. hybrida, and reduced P. intermedia as a va- riety of the latter, P. hybrida var. intermedia (C. А. Meyer) Krylov, in addition to the typical variety. In the area of West Tienshan (Kirghizia and Xinjiang. China), Trautvetter (1904) enumerated only P. an- omala, treating Р. intermedia as its subspecies. Schipezinsky (1921) basically followed Krylov (1901), recognizing two species, P. anomala (with two varieties, var. anomala and var. nudicarpa) and P. hybrida (with two varieties, var. hybrida and var. intermedia), and the same treatment was adopted for the Flora of the USSR (Schipezinsky, 1937) These two Paeonia species were grouped by him as series 4, Dentatae Kom. (Schipezinsky, 1937: 33, "leaf lobes incised or with dentate margin"). Roots of both species were described as tuberous. In con- trast, Stern (1946) recognized only one species in his monograph of Paeonia, P. anomala, in which two varieties were recognized: variety anomala with glabrous carpels, and variety intermedia with to- mentose carpels. He did not mention roots of this 946: 113) considered P. hybrida Pal- las as an ambiguous name: “It is doubtful what group. Stern ( species Pallas intended by this name and, as it is not possible to discover what he meant, this name of P. hybrida has been omitted." However, in the Flora of Kazakhstan, Gamaulova (1961), following Krylov (1901) and Schipezinsky (1921, 1937), de- scribed two species, and still recognized P. hybrida as a valid name. While working on Paeonia for the Flora Reipub- licae Popularis Sinicae, Pan (1979) found a speci- men from Xinjiang whose roots were basipetally at- Schipezinsky (1937) and Paeonia anomala and P. hybri- tenuate. According to Gamaulova (1961), da have tuberous or fusiform roots, so the plant was described by her as a new species, Р. sinjiangensis . Y. Pan; the plants with tuberous or fusiform roots were treated as P. anomala with two varieties: va- Also from Xinjiang, Dai and Ying (1990) described another new species, P. altaica, which was stated to have flowers larger than P. sinjiangensis and one or two underdeveloped flower buds in addition to the terminal blooming flower. Interestingly, Schmitt (1999) failed to recognize the distinct differences between Paeonia anomala and P intermedia in the root and calyx, and thus treated them as a single species, but still recog- nized P. sinjiangensis and P. altaica as distinct spe- cles. From the above it is clear that Paeonia hybrida, P. intermedia, and P. anomala have been variously treated. With this paper we hope to clarify the bi- ological attributes and variations of the P. anomala complex. OBSERVATIONS AND DISCUSSION To answer how many species were really present in Xinjiang and what biological features they have, 1994) made an expedition to the Tienshan and Altai the first author and coworkers (Hong et al., Mountains, conducting field observations and pop- ulation samplings of Paeonia in 1993. Seven pop- ulations in total were observed, covering those character states previously described for the P. an- omala complex (Table 1). We found there were only two species in Xinjiang that differed distinctly from each other in the root and calyx. One species had the roots basipetally attenuate (carrot-shaped) and sepals mostly caudate (Fig. 2), preferring relatively moist habitats in woods. This was treated as P. sin- Jiangensis. The other had roots tuberous or fusiform and sepals mostly (at least 2) rounded but not cau- date at apex, growing on sunny, shrubby or grassy slopes, or in sparse woods. This latter was recog- nized as Р. anomala. We also found the presence or absence of indumentum on the carpels in this species group to be a polymorphism, i.e., individ- uals with carpels either glabrous or from sparsely to densely pubescent could be found within a single population (Table 1). The indumentum on carpels was demonstrated as quite variable, although it was considered by the previous authors as taxonomi- cally valuable. Therefore, any taxonomic division based on this character, such as that between P. anomala and P. anomala var. nudicarpa (by Huth, 2) « co ` P. anomala and P. anomala var. inter- media (by Stern, 1946), is artificial. Also not ob- served by the previous authors was the clear dif- ferentiation in root and calyx in this group. which are closely correlated with each other (Fig. 2). We also found that in the form with carrot-shaped Volume 91, Number 1 20 Hong & Pan 89 Paeonia anomala Complex Table 1. character states, while those in parentheses indicate the number of individuals observed. The indumentum on carpels is divided more or less arbitrarily into six grades with “О” The populations sampled and their characters in the Paeonia anomala complex in Xinjiang, China (all the vouchers are in PE). The figures before parentheses indicate “1” extremely sparsely hairy, “5” entirely covered with hairs, while 72," “3,” and "4" are in between. ssing totally glabrous. > ехрге Ind umentum Number of non-caudate sepals on carpels 0(2); 101); Leaf color Roots Altai: Mt. Halamaryi, NW of Altay City, 1 Locality and habitat Population D. Y. Hong et al. 3 5(1) e pale green carrot-shaped (3) 200 m, Pop- ulus forest by stream Altai: Xiaodong Gou Valley, NE 00) carrot-shaped (3) pale green m, Populus-Betula-Picea forest at valley bottom of Altay City, 1060 Altai: Habahe County, 1200-1550 m. in Betula forests 4 D. Y. Hong et al. 5 4): 0(2) — 0 T. H. Ying 1006—1022 D. Y. Hong et al. 2 'arrot-shaped (2) fusiform to tuberous (3) 2(1): 3(5) 0(1): 20): 3(5) green . Halamaryi NW of Altay City. 1300 m, S slope, rocky Berberis-Spiraea bushes Altai: Mt. Halamaryi, NW of Altay City. 1000 m. Mt. Altai: green fusiform to tuberous (2) 5 D. Y. Hong et al. 012 sparse bushes Altai: Xiaodong Gou, NE of Altay € green 3(2): 41) 0(1): 5(8) fusiform to tuberous (6) 4 D. Y. Hong et al. 4 ity, 1060 m. W slope, rocky sparse Spiraea-Berberis bushes Tienshan: Yining County, Yining Forest Farm, 1150— 2(4): 3(1) green fusiform to tuberous (5) D. Y. Hong et al. 1 1200 m. N slope. sparse Prunus-Malus forest with Cotoneaster & Spiraea roots, the petals varied greatly in length, from 3.5 to 6.5 em, and in the Altai Mountains some indi- viduals of this form possessed one or two additional underdeveloped flower buds similar to Paeonia veitchii. This was again a polymorphism. and not considered by us to be a stable character, and therefore P. altaica was reduced to a synonym of P. sinjiangensis (Hong et al., 1994). ler examination of about 200 sheets of her- m specimens in the Komarov Institute of Bot- any (LE) in 1999, the first author found that two species could also be distinguished in this complex: one with the roots carrot-shaped and sepals mostly caudate. while the other with the roots tuberous or fusiform. and sepals mostly non-caudate. These bi- ological features match those on plants observed in Xinjiang (Hong et al., 1994). Gamaulova (1961, tab. 2) described roots of Paeonia anomala as fu- siform-thickened, and those of P. hybrida as tuber- ous-thickened. However, roots of P. intermedia (— P. hybrida sensu Schipezinsky and Gamaulova) vary in shape from tuberous consistently to fusi- form, and this variation could be found within pop- ulations or even on the same individual, whereas in P anomala neither fusiform-thickening nor tu- berous-thickening were found. Contradictions exist between our natural obser- vations (Hong et al., 1994) and earlier descriptions by Schipezinsky (1937) and Gamaulova (1961). Stern's (1946) treatment of this complex as one spe- cies with two varieties (Р. anomala and P. anomala var. intermedia) is also at variance. For a clear tax- onomic revision and correct nomenclature, one re- maining issue ought to be resolved: the identity of Paeonia anomala, P. hybrida, and P. intermedia. The type specimen of P. anomala, with two visible sepals both caudate and relatively thin leaves. is perfectly consistent with the form with carrot- shaped roots. According to the type specimen at LINN [Siberia, no. 692. 3], fi examination of over 350 sheets of exsiccatae, Р. eld observation, and sinjiangensis and P. altaica should be treated as synonyms of P. anomala; the descriptions of roots of P. anomala by Schipezinsky (1937) and Gamau- lova (1961) are erroneous. The type of P. hybrida (at BM), on which Pallas's (1789) tab. 86 was ap- parently based, has leaf segments 3—4 mm in width. and at least more than 100 in number, and is thus within t of variation in Р. tenuifolia L. (Hong & Zhou. 2003) "Therefore. P. hybrida Pallas should be treated as a synonym of P. tenuifolia, as Anderson (1818) suggested nearly 200 vears ago. The type specimen of P. intermedia at K possesses tuberous roots, mostly non-caudate sepals. and rel- atively thick and narrow leaf segments. The plants 90 Annals of the Missouri Botanical Garden that have tuberous or fusiform roots, are distributed widely from the Altai to Tadzhikistan and Uzbeki- stan, and were named P. hybrida by Schipezinsky (1921, this type specimen very well. 1937) and Gamaulova (1961) correspond to Stern’s (1946) statement concerning the name of Paeonia hybrida is not correct, confusing P. inter- media with P. anomala. Pan's (1979) treatment of the P. anomala complex (a new species, P. sinjian- gensis, and P. anomala with var. intermedia) is also unjustifiable. Although Hong and his coworkers (Hong et al., 1994) properly described two species and clearly stated their morphological and ecolog- ical differences for this group, the nomenclature they used is misapplied. The Paeonia anomala complex comprises two species: Р. anomala and P. intermedia. Paeonia emodi, which was treated as a variety of P anomala by Huth (1892), remains a distinct species, with leaves ternate, without bristles on the upper sur- face, leaflets much less segmented, segments no more than 40 in number and 1.54 cm wide. car- pels usually two, less frequently one, very rarely three, and flowers white. Paeonia veitchii Lynch was described from Sichuan Province, China (Lynch, 1890), and was said (Stern, 1946; 1979) to differ from the P. anomala group in having Pan, several flowers instead of a single flower on a stem. According to our observation, one or two under- developed flower buds sometimes also exist in ad- dition to one normal and terminal flower in P. an- omala, while P. solitary (P. veitchii var. uniflora), although it has З veitchii sometimes has flowers or 4 flowers or 1 to 3 underdeveloped flower buds in addition to the terminal flower on a stem. Thus, P. veitchii was reduced to a subspecies of P an- omala (Hong et al., 2001) KEY TO THE PAEONIA ANOMALA COMPLEX la. Roots carrot-shaped, never tuberous or fusiform: sepals all or mostly caudate, less frequently one or very rarely two P. anomala 2a. Flowers s r 2 underdevel- oped flower buds present in SN to ler- = = 1 5 ш = E ш minal flower on а stem la. P. anomala n ve а Flowers usually 2 to 4 on a stem or 1 to 3 underdeveloped flower buds present in ate rarely flowers sol- = dition to terminal flower, itary yn a stem .P animals 1 veitchii Ib. Roots fusiform or tuberous; inner 2 te rounded but non-caudate at apex 2. sepals | P. intermedia TAXONOMIC TREATMENT 1. Paeonia anomala l.. Mant. 2: 247. 1771. TYPE: Siberia, no. 692.3 (holotype, LINN!). Figure 1A. Paeonia laciniata к аз, Fl. Ross. 1(2): 93, tab. 85, sub P sibirica, 1789. TYPE: the illustration (tab.) cited the аа (lectotype, designated here!). Paeonia sibirica Pallas, Fl. Ross. 1(2): tab. 85. 1789. Pallas's (1789) illustration (tab.) 85 has only one name, P sibirica, and apparently he used this il- lustration also for P. laciniata because one sees "Paeonia laciniata Tab. LXXXV" on p. 93. Paeon- ia laciniata was described in detail by Pallas, but no description of P. sibirica was given by him. Perennials: tap roots = 50 em long. carrot- shaped, basipetally attenuate, up to 2 cm diam., lateral roots slender, neither tuberous nor fusiform. Leaves biternate; leaflets finely segmented; lower leaves with segments 70 to 100 in number, 8-32 mm in width. Flowers solitary or 2 to 4 on a stem, often only terminal one fully developed and bloom- ing; sepals 3 to 5, mostly caudate at apex, rarely or very occasionally 2 non-caudate; carpels 2 to 5, from glabrous to densely tomentose. The species prefers relatively moist habitats, growing in forests, on the edges of forests, or rarely in bushes. It is found at altitudes from 1100 to 3870 m. This species is widely distributed from central China to the Kola Peninsula of Russia via Siberia and Central Asia. The species is of two allopatric subspecies, with the typical subspecies distributed northwest of the Gobi Desert, while subspecies veit- chii is in China southeast of the Gobi Desert (Fig. 1а. Paeonia anomala Ll anomala Paeonia sinjiangensis K. Y. Fl. Reipubl. Popularis Sin, 27: 603, fig. 12. ao тү! PE: China. Xinjiang: Habahe County, Larix forests, 1973, Y. R. Ling 1141 (holotype, PE!). ia altaica K. M. Da „ H. Ying, Bull. Bot. Res. arbin) 10(4): 33, e n TYPE: China. Xin- Tank prre County, Wuzliti ti, 1550 m. in forests, 10 June : Н. Ying 1007 (holotype, SHMU; isolype, pu The typical subspecies is usually found in de- ciduous or conifer forests, in valleys, less frequent- ly in meadows, at altitudes from 1100 to 2200 m. Apparently it prefers relatively moist habitats. This subspecies is of very wide distribution: northeast Kazakhstan, northern Mongolia, from the Altai and Baikal to the Kola Peninsula in Russia, and in the Altai and adjacent regions in Xinjiang, China (Fig. 3). Additional и 5 CHIN lay, J. R. Xu s.n. Ads. . Halamaryi, "n Population No. ; ey, T. H. Ying 7 55 (PE, SHM Cui 86624 (XJNU). Emin: Shiyue онак. Wuerkehe'er, ver ag a Hong « Hong & Pan Volume 91, Number 1 2004 Paeonia anomala Complex Q E | 4 pou лга А МАШ 2 oreet e PETRO > бүтүү! 1,» „ py | = ра BABU пнев! муре 722 ч?” 9477 |, ape cw 14811 LOMO] 1F [әче our jo yuouadır[ua ou st gy3 2oddn үе posut our (М) ләләр ^V 77) P pawon q 7— (NE) SEd »puq J 9 (NNTP »ppuoun а ү suounaads ad&jo|oq vuong р олпат Annals of the Missouri Botanical Garden Figure 2. Diagnostic features of Paeonia anomala (A, В) and P. intermedia (С, D). Tuberous or fusiform roots are correlated with non-caudate sepals, while carrot-shaped roots correlated with usually caudate sepals. right bank of Telikete River, Xinjiang Branch Acad. Sinica 012 (ХЈВІ). Fuhai: Ahebai, glades, s. coll. 3867 (XJBD: Daqiao Forest Farm, T. H. Ying 1011 (PE, SHMU). Fuyun: . W. Chang 11356 (ХЈВІ). Habahe: e ae Z. M. Mao 7 75 (XJBD; Tielieke, fea Valley keli et al. Xingfu Town nship, ey, n ju 7305 (XJBD. Qinghe: ш 1 85 ng Valley, Integr. avo 11829 (XJBI). Tacheng: Mt. Ba'erleike, X. Y Li (SHI). Toli: layer Gold Mine, №. А. Cui 091 е. Mt. Zayier, N slopes, Integr. Exped. (Biology) pa (XJBI). Wenquan: Nerxiaoto Gou, А M. Zhang 90-346 (XJNU). Yumin: Ku'erzhai, Liu & Shen 8308 (XIB I). da TAN. EE Province: Altai, near Altaiskay Station, Mt. Narymsky ео gorge slopes, Таду 92 LE). E Altai: Mt. Ivanovs sky Range, near Gorno- Ulbinki, J. Rohderi s.n. p E). Ustkamenogorsk region: Mt. Altai, Kunduzda River valley, е 360 (LE); Mt. Kalbinskie, Seryal Day River у | Rezni- chenko 49 (LE); Mt. Kalbinskie, Ulansky, near Fée ren by Tainta River, P. Yurchenko s.n. (LE); Mt. Narymsk y, upper U rkar River, V. Reznichenko 58 (LE); near U Ibinsky (LE); Urunhaika M mouth, S banks A. Sedelnikov s.n. (LE). Zaisan region: Mt. Altai, Ulkunc 1 K River alle ғу, upper Kaldzhir Riv- er, V. Reznichenko 106 (LE); Markakul Lake, B. Keller s.n. — — > — Se T LE); Mt. Narymsk y Range, Ak-tui River valley, D. Ya- же lev 69 (LE); nichenko s.n. (LE); Mt. S ley, B. Keller s.n. (LE). MONG OL IA. Ulan Bator: 10 mi. E of Ulan Bator, P. Morgan 23 (K). Gloucester: Mongo- Пее nines с Exped. М. P. Price 293 (К). North . Saldzharsky Pass, E. Klements 11a (LE). e Region: Altai: between Eu po balina, mtn. slopes, E. K. Klements ad Saadak-lary, Vereis 296 (LE). vennitehny, near Nizhne-U n. S. Kolomoitseva 236 (LE). Buryato-Mongolian Autonomous State: Baikal, m a s.n. (LE); Zabaikalje, upper Toreika River, 130 km W of Troitskosavsk, P. Mihno s.n (1 Eubinsk R : West Sayan, Mt. Alan Range, Go- lets Ta Ka I. Krasnoborov 6765 (LE); near Buiba St tion, by river, ed & Sannikova s.n. (LE). Tuva: Tandinsky PEN Mt. Tannudla Range, near Shurmak, V Hanminchun an (LE); Ulug-Hemsky distr., Uiuksky Mtn. Ran ajan-Kol River valley, Orto- Hem tributary, Lomonosova & h 'anova 89 (LE). Irkutsk Region: Irkutsk Prov.: Lena River valley, upper Culm “ay Г Ganda. LE). lala distr.: near r Bazheevskoye, N. Mal- tsev 79 (LE). Nizhneudinsky distr.: by Angara River, op- posite Yandov, 5. in s.n. (LE); Mt. Hariba-bim Range, Angara River valley, Karpovskoye, A. Korovkin 290 (LE); by Oka River, А. "Krishtofow ihe s.n. (LE); near ~ — — = Volume 91, Number 1 2004 Hong & Pan Paeonia anomala Complex Vidimka, Vidim River, S. Kucherovskaya 282 (LE). Tunkin distr.: Mt. Sayany, near Nylova Pustyn, Y. Komarov s.n. (LE): Nizhnaya Tunguska valley. near Nizhnaya Korelina, S. Kokulin 254 ae? Tunkinsky distr.: Smirnov s.n. (LE). Verholensky distr: shin ada 179 (LE): Harat, б. (LE); Lena River & Kirenga valley. Toptykar 67 (LE); Tutura. P. Alexandrov 219 (LE). Kola lin- sula: Ponoi River, left bank, 8 km from its mouth. E. Chernov 3 (LE); Ponoi River, near its mouth, V. Orlova ) (LE). Krasnoyarsk Belsk distr.: near Al- shat. J. Kuznetsov 4661 (LE); Chadobets. G. Borowikow s.n. (LE): Rybnoe. Dranitsyn & Kochubei s.n. (LE). Enisei Prov., Achinsk distr.: near Ingol Lake, A. Suhareva s.n. (LE); near Maloye ree M. Бино а s.n. (LE); Bolshe- Uluiskaya, near Bobrovki, J. Kuznetsov 53 (LE): тг River Valley, J. Тиен 247 (LE): Katanga, G. Bo- rovikov s.n. (LE): Vvezzhi Log. A. Tugarinor s.n. LE. Kansk distr.: Gutar River valley, by Kamenka tributary. W. Troitsky s.n. (LE): near Kansk. A. Shliahtin s.n. (LE): Rybnaya River valley, Perovskoye. J. Kuznetsov 193 (LE): near Nazimovskoye. Z. Evseeva 4406 (LE). Krasnoyarsk Prov.: Chunia River, right bank opposite mouth of Mutorai River, A. Rubin s.n. (LE); Hakassia, Shirinsky distr. Bol- shaya liusa in valley, с: slopes, Polozhii & iig sova s.n. (LE): near Krasnoyarsk, by Mohovaya Rive т skaya & М. "Mishin 104 (LE): Minusinsk. Bols bos vikul Lake. K. Golubeva et al. s.n. (LE): N izhnava Tun- dr River valley, near mouth of Hurkakit River, Rubin & Maskil s.n. (L F): between Salba & Grigorievka, P. lov s.n. (LE): Shushensky distr. near Tanzyben, by Black 'anzyben River, Kuminova & Alexeeva s.n. (LE): near So- rokino Station. N slo )pes. Серыя s.n. (LE). Nov osiber- ia Region: Altai: Elikmanar, upper Karakol River. V. Ver- eshagin 345 (LE). Kuznetsky Alatau: Kondoma iw va +A Kazany Mine. Mt. Kyon, B. Klopotor s.n. (LE): up- per Sary-C humysh River, near Ulus Munai, A. Vydrin s.n. (LE); Tom River valley, ee owed taiga, B. Klopotov s.n. (LE). Mariinsky Prov. j. Talitskiebelky, to Kazanda, Pobe б 697 mE . Lebed River ime Кгу- > СУ i" AC S т = ~. = = - ES ч — ` (LE } i Сб & Chet River valley, P. River valley, a & Sabitov s.n. distr, near Mirny, Lashinsky & Йа ыйа 973 Omsk парон "Tobolsk рилсе: Berezov distr.. ry of Severnaya Sosva. s. re s.n. (LE): pibe River valley, E. Vislouh 73 (LE). Ural: Lyapin tributary, upper Manja River, y гы! have 100 (LE); Severnaya Sosva River valley, Lyapin tributary, ee Khulga River, B. Gorodkov 512 (LE). Mar- Beket River valley, Kolsonskoye. V. Kuznet- . Tobolsk Province: Bere; zovsky distr., Bere- Rozhdestvensky s.n. (LE): Tarsky distr., Rybinsk, near Pustynnoye, V. Varentsov s.n. (LE). Yakutsk | ppc State: Yakutia, Suntarsky distr.. upper Vil- | River, Tuoikhaia, by Chona River, 150/22 (LE). lb. Paeonia anomala subsp. veitehii (Lynch) D. Y. Hong & К. Y. Pan, Novon 11: 317. 2001. Paeonia veitchit Lynch, Chron. 46: 2 tab. 1. Gard. J. Kildushewsky ser. 3. 1909. TYPE: China. W. Sichuan: Tatien-lu (Kangding), 8—10.000 ft.. E. H. Wil- son (for James Veitch & Sons) 3034 (holotype. K!, photo, PE!). Аи = beresowskit ا ا‎ Bot. Mater. Gerb. Glavn. Bot. Sada RSFSR 2: 15 Jan. 1921. Lx shit (Komaroy) hipezinsk y. SR 2: 46. 26 Mar. Paeonia veit- we var. Mater. Gerb. 6 Gia). Bot. Sada RS 1921. China Oecid. a huan (Sichuan). Sun-pan-tin (бо gpan). Guichua.” 9 June 1894. M Be resowskii s.n. (lectotype, designated here. LE seen). Paeonia veitchii var. 5 = ex Cox) Stern, J. aay Soc. 69: 1943. Paeonia ioodwardii Stapf e Pl. Int md Farrer 43. 1930. TYPE: 1 Zone (Chuoni). K. Farrer 67 (holo- type. E not seen). Paeonia veitchii var. leioc arpa W. J. Wang & S. Н FI. Reipubl. Mi ue Sin. PA 603. 1979. TYPE: me Sichuan: Jinchuan. Kasa Township. Yin- changgou, forests by stream, 2700 m. 26 Apr. 1958. 77248 Gel PE”. . eile ‘chit var. uniflora К. Y. pularis Sin, 27: 603. Gaze, Xiongjiling. : 3000 m. mtn. summit. June 1974. فا‎ Xizang Exped. Veget. Erin 034 PE!) по! Y Roy. ^ ~ 1 ‚ Wang, Рап, FI. ‘publ. Po- (holotype. This subspecies, though distantly isolated from Paeonia anomala subsp. anomala by the huge de- serts of the Gobi, still very much resembles the latter. The only remarkable difference between them is the number of flowers on a stem. Paeonia anomala subsp. veitchii usually possesses 2 to 4 blooming flowers in addition to up to 2 underde- veloped flower buds. Very rarely only the terminal flower blooms in addition to up to 3 flower buds. Paeonia anomala subsp. anomala possesses only a single terminal blooming flower, without or infre- quently with 1 to 3 underdeveloped flower buds. The subspecies is widely distributed in China: southeastern and central Gansu, southern Ningxia, the Qinling Range of Shaanxi, and the eastern extreme eastern Qinghai, Shanxi, western Sichuan, of Xizang (Tibet) (Fig. 3). Like subspecies anom- ala, Paeonia anomala subsp. veitchii prefers rela- tively moist habitats, growing in forests, grasses on the edges of forests, bushes, or subalpine and al- pine meadows with shrubs, at altitudes from 1800 to 3870 m. Phenology. April to early June, and fruits in August and Sep- The subspecies flowers from late tember. Chromosomes. 2n = 10 (Hong et al., 1988). Additional Tp, examined. CHINA. Gansu: Di- ngxi, s. coll. s.n. (CPB). Hezheng: Xinzuang. Gansu Herbs сар. s.n. (NWTC). E da Mt. Dalinke, Q. R. We Ш . Jiangcha: E У à Suet Kangle: Mt. . Lianhua, M. S. Yan 17 55 (NWTC). Kangxian: s. loco, 94 Annals of the Missouri Botanical Garden x PN \ 0 ¿NO Pd N pt ‘S X БА 2 60° e hi m D MEE nm a e 30° > ө 130° > ө ө 50° e e ® 40° e e e * e е е gon 222 227 * * 72 40° * * Tk * 4 * Ру да. о оо 120° 9069 о 50° OX? о Соо О 30° y о C2 со 8 O Oo? la 6 ~D O 60° 70° 80° 90° 100° 110° Figure 3. anomala; white circles: / or several vouchers cited in the text. 5 coll. s. n. uera Lanzhou: M Xinlong, s. coll. 84 С Tiandu, s. 1 s.n. (NWTC). Linxia: near y 834 (РЕ). Longdie: Dacaopo, 1. J. Zhou 708 w Мо: Lujing, Zhongchuan- gou, J. Q. Wang 197 (NWTC); Mt. Luodadoujidela, J. P Wang 15240 (PE); near Mawu, T. P. Wang 4594 (PE); between Lintao and egi Ms dis nos 1689 e 7 Mt. Wutai, Taohe Exped. 3 3 (РЕ); Lamathan, K. S. 604 (PE). Taohe: J a 12829 (PE). Tsaluku to shi men, J. E Rock er (PE). Tianshui: i Xinjia Hsia 5707 (PE).“ : Anyuan, У. He 4316 (PE); Zhucha, Y. Q. He 4915 (PE); ); Jinqiang үш. ip, Mt. Мао- mao, Q. К. Wang 1869 NN). Xiahe: Chingshui, 7. P Wang 6944 (PE); Tangarang. 1 О. К. Wang 7270 (NWTC). Yongdeng: Liane heng, Tulugou, J. L. Bai 8613 (NWTC). Yuzhong: Xinlong, near Venida aowan, s. coll. s.n. (PE). Zhangxian: Сеш. distr., 1 s ^d. 03183 (PE); Mt. Guiqing, 1 od wre 4735 (PE); Hediling, Mt. Kehu, Huanghe Exped. 4655 (P E Shi- chuan Forest Farm, Jinhuacl thi, Lian, Wang et al. 79197 (NWTC). Zhouqu: Taozhou Forest Farm, Jiang & Jin 00386 (PE). a (Chuoni): Kache Forest Farm, Lian & 2o 31 (NW ©); Wanchang, Sangtanagou, Lian & Chen pbi = — — = S © = 2 Е. O; No © f — ' vu = — Dongxia, Yu, Lu, Gu & Li 68 (PE); Mt. Laoye, Liang et Distribution map of the Paeonia anomala species complex. Black c "ircles: Paeonia anomala subsp. Р. anomala subsp. veitchii; stars: P. intermedia. Each site in nis map is supported by a single al. 378 (HNWP); Baoku, Z. H. Zhang et al. 4291 HNWP). Huangyuan: Mt. Banjie, P. C. Meg 8842 (PE). 5 Sanhei Forest Farm. Wang 1065 HNWP); 40 km W of Xining, S. А Zhen 0351 (PE). Huzhu: ae Forest Farm, Guo & He 9012 (HNWP); Qiaotou, Guo & Wang 6717 Р). Jainca: near Angla, Liu & Luo 1040 (NWP) Pedi Maying 3 Ko- ngjiazui, В. Z. Guo 6767 (HNWP). Golog: M Farm, Hongjungou, W. Y. Wang 26818 (HNWP). Menyuan: Semnyi, betw. Lihua & Dalong, Z. Y. Qing 1218 (HNW PE); Semnyi, Zhugusi, Xielong. B. Z. Guo 7396 (HNWP): Semnyi, Hankegou, Cans “Qinghai Exped. 2493 (PE). Minhe: Gushan, Nanxia, I. H. Zhou 2528 (HNW P); ien erbeishan, B. Z. Guo 7010 (HNWP). Tongren: Rong- », В. Z. Guo 10232 (HNWP); Shuangfengxi, B. Z. Guo 10186 (HNWP). Xining [Sining]: Shangwuchuang, К. S. Hao 779 (PE) Xunhua: Mengda Forest Farm, Guo & Wang 25058 (HNWP); Il / WP). Shaanxi: Huxiar Phatinae. ig 2019 (PE). 1 longbei, W. J. Hsia 4539 (PE). М. Та temple, W. Y. Hsia 4571 (PE); Sanchaixia, К. J. Fu 4441 PE): Dadian temple to oo temple, Hong & Zhu PB85065 (PE). Shanxi: Seo ie Wang & Tian 594 (PE). M lage, Xiejiagou, Shanxi Exped. 684 (РЕ). Sichuan: Baox- ing: Raozhi, Nibagou, Hong & Zhong PB82105 (PE): p — — p Volume 91, Number 1 2004 Hong & Pan 95 Paeonia anomala Complex Е Zhonggan. МІ. 1 5 5 459-0246 (PE); Ganvanggou. I. T. () (PE); Dengch- igou. 7. P Tu 4329 (PE): Lianghe a PM ele p & Ren 5572 (PE). Barkam: near Puvajiao. A. Li 71029 (PE); near Kanzhugou. by river V. Li 70441 (PE): Pwermagou. 209 Lumbering Ground. M. Li 70867 (PE): Bwermagou, X. Li 70903 (PE): near Maan: Li 71645 (PE); Dalongjiaogou, X. Li 70685 (PE): near kafeiqiao. A. Li 71196 (PE); Zonggag. X. Li 70719 (PE): Nazhugou. X. Li 71087 (PE). Emei: Jieyindian, J. H. Tu 362 (PE). Dege S. X. Jia 229 (PE). Garze: S bank of Yalongjia River, Y W Cui 4348 (PE). Heishui: Shidiaolou Township. Sich- uan-Econom. 459-1208 (PE): Sandougou, Sichnuan-Econ- om. A59-1001 (PE). Jinchuan: Zosijia. E X. Li 10157 (P r4 Jiulong: Mt. Baitai, between Niupo & Baitai village. s. Ying 3869 (PE). Kangding: Yulingong. Yingbapo. Hu & He e (PE): Yulin Township. — Jiang & 5 (PE): Yulin ee 8 . C. Jiang Taivangshan distr., Hu & He ve (PE): Niugu village. W Sichuan Exped. oe et al. 243 (РЕ); Xinduqiao. W Sichuan Exped. Kuan et al. 732 (PE): Shade distr. M Along. Nanshui-Beidiao Exped. 02912 (PE): Zhonggu village. Dagaigou. W. Sichuan E n d. Kuan et al. 352 (PE): between Kangding & Dawu. P C. Tsoong 5004 (PE). Lixian: Longxi. Yiduo village. Z. He 12559 (PE): Mivaluo. bo, yu, S. Jiang A-7162 (PE): Shuan- Shu: anjingsi, 333 km milestone of € eal highway. Zhang & Zhou 23742 (Pk): Somang. 12 km milestone of Shuanjingsi— Багтай high- . Shang & Zhou 22675 (PE). Ts 5021 (РЕ); 2 hor. by rive jingsi. P X. Li 10100 (PE): F. T. Wang 22930 (P "E . Mafang. Sichuan Econom. (4-59) 2554 (РЕ); Зачем аһа. Sanlonggou, He & {ма 1 I. d. 138 (PE). Mianning: Yejin Township. 5. E Zhu ; Huning distr., Jiexingou, S. Jiang 5708 (PE). ! Ун 6137 (PE). Qianning: S of town, Jiang & Jin 2113 (PE); to Dawu, Zhang & Lang 57 (PE). Sêrtar: 1 er S of Wuneda, S. Jiang 9082 (PE). Songpan: m c S. Jiang A-7275 TE Mont. Occid. H. Smith (PE). Tianquan: Mt. Erlong. Niaoniujingou. around 2 Hu & He 10088 (P E). we nec huan: Wolong Town- D ru ed 459-2290 (PE): Wolong Nature Zalang. 2 & fin Ph 85019 (PE). Xiao- jin: Xihe VR Niuchanggou. Zha | Ren 6322 (PE): Luobigou, bel N Zhang к qe 6082 (PE). Ya- Mt. Ermin, Sichuan Plants Collection 0905 (РЕ); Tanjiao to Malangeuo, J. S. Ying 3113 (PE). Zamtang: e. right bank, S. Jiang 8930 (PE); ТО km № of lbangmuda, S. Jiang 8965 5 (PE). Xizang (Tibet): Jomda: Gamtog distr.. Qinghai-Nizang Exped. Vegetation Group 9893 (PE). ship. Resery YG; jiang: Lukesi Temple Lede- A. Meyer, in 1830. Paeonia anom- Paeonia intermedia C. Altaic 2: 277. N bour, Fl. ala var. hybrida f. intermedia (C. Meyer) Trautvetter. Enum. PI. Songor. 88. 1800. Paeonia hybrida var. intermedia (C. A. Meyer) 1: 47. 19 Ol. Paeonia an- . A. Meyer) Traut- Tidi Imp. S. Pe- 1904. Paeonia Meyer) O. & Krylov. Fl. Altava omala subsp. intermedia (C Fedtschenko. Sada 23: 351. intermedia (C. A. Beih. Bot. Centralbl. in B. Bot. anomala var. B. Fedtsch.. veller, tersburg. 18: 216. Ledebour TYPE: the Altar Mountains, (lectotype, designated here, K!). 1905. Paeonia gon N. he 1zchoweli. Notul. Syst. Geogr. Inst. Bot. l'iphlis. f asc. 21: 17. 1959. TY PE: Ge orgia. ipo between Igoeti and Lamitskali, 16 May 195 TBI! tschoweli s.n. (holotype. ). Paeonta interme subsp. sig ceu * à Ovchinni- ov. dH Tadzhikistanskot SSR. -10 & 531. 1975. nL Padzhikistan: dec cod ipii jugi Hissarci ic. ad ripam m fl. Maichura. 5 km ab ostio. 2500 m. 12.06.197 1. Glebowa 6 (holo- pe. T AD not pre qd The species has long been identified as Paeonta anomala Т 197 d treated as a variety of P 1901). i schenko, 1905: 1946: subspecies of P. anomala (Trautvetter, maulova, Pan, 1979), a 1904). o even as a form of P. anomala var. hybrida (Vraut- vetter, 1860). These diverse and erroneous assign- ments were made (1) the identity of P. anomala, P. intermedia, and P. hybrida was not pre- viously clear; (2) previous authors emphasized the taxonomic value of indumentum on carpels; and (3) Ps because: the characters of roots and calyces and their cor- relation were not considered. Examination of the types of the three taxa and extensive observations on herbarium specimens and natural populations Table 1) independent species, distinctly differing from X show that P. intermedia C. A. Meyer is an — anomala by its lateral roots tuberous to long-fusi- form, sepals mostly (at least 2) rounded at apex but not caudate. Also different from P. anomala, P. in- termedia prefers relatively sunny and dry habitats. Carpels vary from | to 5 in number. and from gla- Ovchinnikov (1975) properly treated the peony in Tadzhikistan as P brous to densely tomentose. intermedia, but his description of the new subspe- cies pamiroalaica is unjustifiable. His description and our extensive observations have not revealed any significant difference. This species is widely distributed in northern Xinjiang. China (south to the Tienshan). Kazakhs- tan, Kirghizia. Tadzhikistan, Uzbekistan. Altai of Russia. One small isolated population was found in Georgia (Hong & Zhou, 2003) (Fig. 3). Ht meadows. and the grows in grassy and shrubby slopes. steppes. or in sparse woods. at altitudes from 900 to 3250 m. Phenology. to late June and fruits from August to September. 10 (Hong et al.. — This species flowers from late May Chromosomes. 2n unpub- lished). Additional specimens examined. CHINA. "pr Altai Mt.: Acad. Sinica Xinjiang Integr. Exped. 1065 Annals of the Missouri Botanical Garden (PE). Altay: Qiao’ati, G. L. Zhu et al. 6325 (PE): to Hal- amaryi, Acad. Sinica 2917 M dos 10227 (PE); Klasu Gou, R. C. Ching 2433 axiong Gou, Т Y. Chou 652100 (XJBI); " B D. Y. Hong et al. Popu- lation No. 2 (PE); Xia = Gou, D. Y. Hong et al. Pop- ulation No. 4 (PE). Bark dpi s. coll. 780 (XJBI). Fuhai: Fuhai Forest Farm, C. L. Zhu 5665 (PE); № slopes, T. Y. Chou et al. 652150 (PE. XJBI). Fukang: Tienci, Shi- men, D. Y. Hong et al. 0190 (PE). Fuyun: Mica No. 3 Mine, /ntegr. Exped. s.n. (XJBI); Mica No. 4 Mine, /ntegr. Exped. s.n. C . Habahe: Tielieke, Kelimu 10171 (XJBD; Wuzliti, Kelimu a JBI). Hoboksar: Songshu Gou s. coll. 7385 31); Mt. Qiaganebo, /ntegr. Sh pi 10563 (PE, XJBI). Huoc e Aksu Commune, Y. R. Lin 74848 (PE); 5 Acad. Sinica Exped. 10402 (PE); i, Yeguolin, W. Chang et al. 3316 (XJBD: Xiaoxigou, А. К. N. Zhu 10414 (PE, XJBI). Mori: Nangou, S. L. Chen 0040 (PE). Qapqal: 58 km e to Zhaosu, Inst. Biology and А к 66 (XJBI). Она: Biliuhe, J. С. Zhao 82-4288 U); Kuangou, Biology 81-8381574 (ХЈВІ); Tangf angen Forest Farm, /ntegr. ] ) iejin Commune, s. Q. Lin 34 (XJBI). Tacheng: Mt. “Bae rleike, N slope, /nteg. Exped. 1200 (XJBI). Toli: Mt. Albakzin, е “т К. C. Kuan 2611 (PE, XJBI). 5 = nsha ү. Chou et al, 650392 (РЕ). Yining: Yinin est oe Tis toulong valley, D. Y. Hong et al. Population. | & 092 (PE). Yumin: уч Wild Bada Prunus Nature Reserve, J. C. Zhao 85-293 ‚ XJU). GEORGIA. Kartly: B. Lamis- суры A ipei Hong & Zhou H99033 (PE). KAZAKHS- Ili-Pri M ridges of Mt. Kirgizsky Range, upper Su ata River, V. Golos-Kokov s.n. (LE); Is- syk-Kul Province, Santash, iesu garden, in & jus Wit s.n. (LE); Semirec 'hensk Province, N slop — >: as = А И, j co Сот ‚ (LE); SW Songorian д Mt. Emelsky E); Taldy-Kurgan Province, Mt. ius -Altyn- Emel pass to road rkent, Grubov & Luboarsky 0 (LE); Zai- liisky Alatau, Usun-Kargali River Valley, W Sukonnaya Fabrika, Y. Goloskokov (1 E). E. Kazakhstan: Mt. Kalbin- ok-tau, near oubinskle е А. Yunatov | distr, Kusto istr., near Kender- Sary- Ozek-Dzha & Borsowa 52 (L sk Prov., Sonnet Alatau, S of Lepsinsk, R. Rozhevits 55 (LE); W ridges of Songorian Alatau, Koksu Gorge, near Koksuis V. Goloskokov (LE); Tarbagatai Chenarak i маа & Buhaeva 298 (LE); Mt eep gorge from the pass, Schipczinsky 5 Kazakhstan: Mt. Chatkalsky Range, Chat- kal Valley, Mt. Piazak, O. Knorring 60 (LE); Prichuisky кү, Sulunger Gorge, V. 5 0 ie 66 (LE KIRGHIZ Dzhalalabad Province aE Kuren distr., Mt. Range, S slopes, near AK-Terek, Le- bedeva & Faleeva s.n. (LE); Kara- Alma, Toguz (Bulak), N slopes, Jie einig s.n. ; Tianshan, Mt. d Range, Mt. Elchin- Ustriuk, Kizilbulak Gorge, s. coll. s E E) E rgana Province: Kirgizsky Alatau, Makbal Gorge, Ty-Karyn River, б. /golkin 97 (L E); Oshsky distr., Ak-Bura valley, Mt. Kanida Pass, O. Knorring 25 (LE). slope, A. Savenkova 29 (LE). RUSSIA. Altai: Alex Shrenk s.n. (LE); Altai, Barnaul Province, between Kalmytskie Музу & Mahanova, P. Krylov s.n. (LE). TADZHIKISTAN. Gissarsky Mountain Range: Buhara, Gissar, Mtn. Zar kamar Pass, V. Lipsky 1697 (LE). Darvazsky Mtn. lur. N slopes, left bank of Zarako River, cai Havdak, Shinova & Kinzikaeva 1550 (LE). Imam-Askari, Mt. W of Darai-Imam, Bochantsev & Egorova 1014 (L " Peter I Range, N slopes, Nazarak Valley, upper reaches, 15 km S of Tadzhikabad, Ladygina, Ikonnikov & Fridman 1337 (LE). Vahshsky Mtn. Range, Mt. Hozreshi-Sho, above Sary-Hosor, Sidorenko 185 (LE). Varzob Valley, right bank of Maihur Gorge, NE slope, Stepanenko & С Кини 2514 1 E); Mountain pass between rivers Hava ts E): upper Horanc hon, ad ве 351 (LE). UZBEKISTAN. Tashkent Provine Bolshoy Chimgan, Z. von Minkwitz 1190 (LE); Bolshoy Chimgan, N stony slopes, V Bochantsev 547 (LE). Literature Cited Anderson, G. 1818. A monograph of the genus Paeonia. rans. Linn. Soc. o 12(1): 248-283. Dai, К. M & T. Н. Ying. 1990. A new species of the genus Paeonia from Chins. Bull. Bot. Res. (Harbin) 1004): 33— к” Candolle, A. P. 1818. ne Vegetabilis Systema Na- urale 1: 386—394. Par . 1824. P verme Systematis Naturalis Regni Ve- getabilis 1: 56-66. Pa Кейс 8 B. A. 19 "n p pop Tyan-shanya 3-105. Se Hort. Petrop. 23(2): 351—353. 1 0 A. P. 1961. Paeonia. Pp. 12— түз М. У. Рау- ov ae Fiora of Kazakhstan, 4. Tab. 2. Alma-Ata. Hong, D. Y. ‚ L. Zhou. 2003. Paeonia (Paeoniaceae) in ЕА 'asus. Bot. J. Linn. Soc. 143: 135-150 ‚ Z. X. Zhang & X. Y. Zhu. 1988. Studies on the genus Paeonia (1)—Report of karyotypes of some wild es in € p Acta Phytotax. Sin. 26(1): 33-34 . Y. Pan 1. 1994. fige, in Xinjiang, China. Ас ta Phy Sin. 32(4): 349-355 ‚ К. Y. Pan & N. J. Turland. 2001. (ner an- omala Eu veitchii (Paeoniaceae), a new combination. 15-318 speci L. 1802. Monographie Gattung Paeonia. En- glers Bot. Jahrb. 14: 276 Krylov, P. 1901. Flora yid Altai 1: 47. Tomsk. F. 1842. Flora Rossica 1: 74. Stuttgart. . 1771. i enn Plantarum. 247. Holmiae. Lynch, R. 1. 1890. А new classification of the genus Марав J. y Hort. Soc. 12: 42 er, C. A. 1830. Paeonia. Pp. 276-279 in C. F. Led- about (editor), Flora Altaica. Berolini d 5 R. 1975. Flora Tadzhikistanskoi SSR 4: 6 0 & 531. I edebour, C. RUN P. s. 1789. Flora Rossica 1(2): 92-95, tab. 84-87. St. Petersburg. Pan, K. Y. 1979. Paeonia. Іп: W. T. Wang (editor), Flora Reipublicae ee Sinicae 27: 37—59. Science Press, Beijir Schipe кзы . 1921. Synopsis of the E nus Paeonia. Not. is en Hort. Bot. Petrop. 2(11-12): 41-47. 7. Paeonia. In: V. L. Komarov (editor), Flora б; 35. Editio Akademii Scientiarum URSS URSS 7 кии 1. Schmitt, E. 1999, Les Pivoines. Étude systematique du enre рар L. Plantes de Montagne 191: 574—583. Stern, F. C. 1946. A Study of the Genus Paeonia. Royal Horticultural Society, London. Trautvetter, E. R. 1860. Enumeratio plantarum songori- Volume 91, Number 1 2004 Hong & Pan Paeonia anomala Complex Alex Schrenk annis 1840-1843 collectar- . 33: 87-88. Fedtschenko (editor), Petrop 23 carum, a Dr. um. are Soc. Nat. Mose 904. ا‎ In: B. Flora a the Tianshan TU Hort. APPENDIX 1 INDEX TO ENSICCATAE EXAMINED The figures in dde ses represent: la = Paeonia an- omala sna ансат: 1b = Р реи subsp. veitchii: 2 = P interm ~ х Acad. Sinica Bot. Inst. Exped. Xinjiang 1803 (РЕ. XJBI) (2): Acad. Sinica Exped. 10402 (PE) (2): Acad. Sin- ica Xinjiang Exped. 2496 (PE, XJBI) (la). 10227 (PE) : Acad. Sinica Xinjiang deus Exped. 10657 (PE) (2): es College 1051 (PE) Каа & Kashenko s.n. (LE) (2): Alexandros, P. 17 zn (LE) ) (La). 219 (LE) (1а): Alex Shrenk s.n. (LE) Bai. J. L. 8613 (NW IC) (Ib): Beresowskii. M. (LE) lb, type of Paeonia beresowskii Komarov); Biology Exped. 79250198 (XJBI) (2). 81 "8330478 (XIBI) (2): Boc dra am V. 547 (LE) (2): Bochantsev € Egorova 1014 (LE) (2): Borovikov. G. s.n. г ) (la): 1 el al. s.n. CS í Jal Chang. 56 (XJBI) i 3316 (X]BD (2). "e (XJBI) A re (ХЈВІ) (la): Chen & Ju s.n. (NW I 1b): 0040 (PE) (2); Cherepnin s.n. (LE) (la): С hing, К. СЕ (PE) (la). А (PE) (2), gts (PE) (1а); А 652100 (XJBI) (2): Chou. et al. eee (PE) (2), 650868 (PE. XJBI) (2). EN 50 (PE. XJBD (2): Cui. N. R. 091 (XJNU) (la), 86624 (XJNU) (la): Cui, Y. W. 4348 — 10255 1803 (PE) (1b). MN (PE) (1b). Dolenko, G. 108 (LE) (la): Dranitsyn & Kochubei s.n. (LE) (1a). Ermolajeva, M. s.n. (LE) (1a); Evseeva, Z. 4406 (LE) (la). Fang, M. P. 4213 (LE) (la), 6037 (LE) (la); Farrer, 67 (E) (Ib. type of Paeonia O Stapf ex Cox): Fedorov & Iljina 72 (L " (2). 128 (LE) (2); Fu. K. J. 4441 (PE) (1b); Fu. T. K. 834 (PE) ( na , in. S. s.n. (LE) (la): Gansu Herbs (NWTC) (Ib): Genina, S. s.n. (LE) (la): Gansu-Qinghai Exped. 2493 (PE) (1b); Goloskokov. V. s.n. (LE) (2): Gon- tscharow. N. 2051 (LE) (2): € a B. 512 (LE) (la): »rigoriev & Buhaeva 298 (LE) (2): Grubov & Luboarsky 229 (LE) (2): Guo, В. 7. 5 E (HNWP) (1b). 7010 (HNWP) (Ib), 7396 (HNWP) (Ib). 10186 (HNWP) (1b), 10232 (HNWP) (1b): Guo & He 9012 (HNWP) (Ib); Guo & Wang 6008 (HNWP) (Ib). 6717 (HNWP) (1b), 25058 (HNWP) (1b); Guo & Yang 9608 (HNWP) 08) 5 V. 515 (LE) (la): Hao. S. 770 (PE) | (0. 4516 (PE) (Ib). 8 75 ir 19 He, 7 : He & Zhou 13338 (PE) (1b): Hong p t al. Тоша 2 (РЕ) (2), pation 3 (PE) (la E. | (PE) (2). deba ie 5 (PE) (la). Population | (PE) (2). TER bein (P E (2 2); л 5 Zhong PB82105 (PE) (1b): Hong D. : Zhu vA (PE) (Ib). PB85040 (PE) 1155 ias (PE) (1b): « He H95034 (А. CAS, K. MO, PE. L Zhou H99033 (PE) (2): Group s.n. ~ . 10463 РЕ) 11173 (РЕ) (1Ь); Hiang, L uo & Time 7 (30 (PE) о, G. 97 (LE) (2): dede & Fridman 892 (LE) (1b). ) (La): Integr. Exped. 630 (XJBI) (1a). 10563 (РЕ. XJBI) (2). Integr. Inst. Hydrol. (2); Iljin. V. s.n. A 1200 (JBI) ( ). 7305 (XJBI) (la). 11829 (XJBI) s.n. (XJBI) (2): Pedol & Biol. Res. 5788 (XJBD (2). Jia, 5. E т (PE) (1b); Jiang & Jin 00386 (PE) (1b). ): Jiang & Xiong 34242 (PE) (1b). 34319 (PE ) (1b). Aid (PE) (1b): Jiang. S. 5768 (PE) (1b), 8874 . 8930 (PE) (Ib). m (PE) (1b). 9082 (PE 4 д y ‚ A-7275 (PE) (1b); Jiang. X. C. ‚ 36433 (PE) (1b). 2}, _ 10369 (AJBD ( 2). 10614 (TBI) (2. type of Paeonia majko N. shewsky. I. 150/22 (LE) (la); Kleme 175 E. 4d (L E lla (LE) (la): ky pa B. 113 (LE) ( iced О. E) (2). 66 (LE) Ам (LE) (1а); cia и S. 236 (LE) (1a); Котагоу, V. s.n. (LE) (la): Korovkin, A. 290 (LE) (la): Krasnoborov. I. 6765 (LE) (la): Krishtofowich, A. s.n. a (LE) (1а); Krylov, P. s.n. (LE) (2); Kuan, К. С. 2611 (PE. XJBD (2), 77309 (PE) (1b); Kuan et al. 243 En er (PE) (1b): Kuch- erovskaya, S. 282 (LE) (1а); Kuminova & Alexeeva s.n. (LE) (la); Kuvajev & Sabitov s.n. (LE) (La): Kuznetsov. I. 53 (LE) (1a). pe y (la). 103 (LE) (la) ШО Kuznetsov, N. 199 (LE) (1a). 1991 (LE) Ladygin 92 S (la), s.n. (LE) (2) Lg et al. 1337 (LE) (2); Lashinsky & 5 973 (LE) (1а): Lebed- eva & Faleeva s.n. (LE) (2); Ledebour, s.n. (К)! 2. type 4 Paeonia intermedia C. ever); Li & Zhu. J. 10414 (PE, XJBD (2): Li 1 В а 10100 (P E) (1b), E (PE) (1b). 10143 (PE) (1b): . 40247 (PE) (Ib). 70441 (PE) (Ib), 70510 (PE) (1b), pret (PE) (1b), 70719 (PE) (Ib). 70867 (PE) (1b). 70903 (PE) (ТЬ). 71029 (PE) (1b). 71087 (PE) (1b), 71196 (PE) (1b). 71645 (PE) (1b), 74842 (PE) (1b), 74936 (PE) (1b). 77248 ( ) (Ib. type ol Paeonia veitchii var. leiocarpa W. T. Wang & S. H. Wang): Li 870147 (SHI) (Та). 87 е (SHI) (2): rie ho 72 (NWTC) (1b): Lian et al. En Licent, P. 1 Chen 31 (NWT (HNWP) (1b). 10) 1255 79179 (NWTC) (Ib): Lin, S. Q. et al. 34 (XJBI) (2): Ling. Y. R. 1141 (PE) (la, type of Paeonia pony nsts К. Y. Pan). 74243 (PE) (2), 74848 (PE) (2). 74849 (PE) (2): Lic 1. 5659 Р a Dor de 6019 (PE) (1b). 61 78 (PE ) (1b): Lips . 1697 (LE) (2): Liu & Luo 1040 (HNW P) Ih: Liu & M 8308 (XJBI) (la): Liu. S. W. 2115 (HN y (1b): Lomonosov a & Ivanova 89 (LE) (la); Long, X. F. é PN | 79 (LE) (la): Mao. Z. M. 10413 (XJBI) (la): Mihno, P. s.n. (LE) (Ja): Minkwitz Z. von 1190 (LE) (2): Morgan, P. 23 (К) (la). ue Beidiao Exped. 02912 (PE) (1b). Pharmac. Exped. 2019 (PE) (1b); Pobedimova 697 (LE) (la) Pohle & Rozhdestvensky s.n. (LE) (1a): Polozhii x Kandasova s.n. (LE) (la): Paniatovskaya s.n. (LE) (2 Price, М. Р. 293 (LE) (1а). Qing, Z. Y. 1218 (HNWP. PE) (1b): Qinghai-Xizang Ex- ped. Veget. س‎ 034 (PE) (1b. type of Paeonia кее var. uniflora ‚ Pan), 9803 (PE) (1b): Qu. J. X. (PE) (1b). Reznic үе у. 49 ( LE) (la). 58 (LE) (la): Reznichen- . 106 (LE) (1a). 360 (LE) (1а), s.n. (LE) (1а); 19 5 (P E ) (1b). 12829 (PE) (ТЬ). 13127 (PE) (1b); Tode. I. s.n. (LE) (la): Rozhevits, R. 55 in. А. s.n. ) i ‚ (LINN) (la. s.n. (2): s. coll. Schipezinsky 125 type of Paeonia бча А Lisa: Sede din: A. 98 Annals of the Missouri Botanical Garden (LE) (1а); S. Gansu Grassland Exped. 681 (NWTC) (1b): бш. vit 684 (PE) (1b): Shanxi Exped. Wang & Tian 594 (PE) (Ib): Shinova & Kinzik aeva 1550 (LE) (2); Shishkin, B. s.n. (LE) (2): * (LE Shukin, S. s.n. (LE) (la); ichua an-Econom. (PE) үе A59-1001 (PE) (1 b A59-1208 (PE) (1b), A59- 2554 (PE) (Ib), Sichuan-Econom. & Ya 838 (PE) (1b): Sichuan Plants Collection 0905 (PE) (1b); Eis 185 (LE) (2); pid M s.n. (LE) (1a): 9 H. 2499 (PE) (1b): Soe A V. 100 (PE) (1b); Sokolov, P. 13a um E) (1а); Song, Z. P. 38544 en (1b). 39107 (PE) (1b); Stepanenko & MM : 2514 (LE) (2); Suhareva, A. s.n. (LE) (1a). Tang, T. 938 (PE) (1b); Teche Exped. 3243 (PE) (1b): Tatarova & p oi 52 (LE) (2); Tolmatschew, I. 247 (LE) (1а); Tomin, M. 67 x E) (la): 5 T. s.n. (LE) (2): Troitsky, M. s.n. (LE) (1а); Tsoong. P. C. 5004 (PE) (1b). 5024 (PE) (1b), 8842 (PE) (1b); Tu, T. H. 362 (PE) (1b). 4329 (PE) (1b); Tugarinov, A. s.n. (LE) (1a); Turezaninow, s.n. (LE) (1a) Varentsov, V. s.n. (LE) (1a); Vereshagin, V. 296 (LE) (la), 345 (LE) (1a); Ve rhovskaya & Mishin 104 (LE) (1a); — E. 73 (LE) (1a); V vdrin, A. s.n. (LE) (1a). Wang, F. T. 22930 (PE) (1b): Wang, J. Q. 197 (NWTC) (1b); W ang, = R. 1869 (NW TC ) (1b). 7034 (NW TC) (1b), 7270 (NWTC) (lb); Wang. 1065 ( а ТОР. al (NWTC) (1а), 15240 (NWTC) (1а), 4594 WTC) (la), 6944 (PE) (1b); Wang, W. Y. 26771 (HNWP) a 26818 (HNW a (1b), 27 060 (HNWP) (1b); Wang, W. d et al. 27347 (HNWP) (1b); Wang & Zhou 19 (HNWP) 1b), 198 A (HNWP) (1b). up Hie th Acad. Sinica 012 (XJBI (1a); Xu, L. 1. (PE) = pe D. 69 (LE) (la); Yan, M. S. 3 (NWTC) (1b); Ying, J. S. 3113 CE) (1b), ان‎ ve) 0, 4582 (PE) (1b), As (PE) (1b); Ying, T. H. vs SHMU (Та), 1001 (PE, SHMU) (Та), 1006 (PE, SHMU la), 1007 (SHMU, PE) (la, type of iuc altaica K. Mr "pt & H. ving, 1008 (PE) (1а), JO (PE) (1b), 6137 (PE) (1b); 1 A. 24 (LE) (2): Yurchenko, P. s.n. (LE) (la Zapriagaev, F. 66 (LE) (2), 351 (I 7 = Zhang, Е M. 90-346 (XFNU) (la); Entes G. Z. 005 (HNWP) (1b); Zhang & Lang 57 (PE) (1b); Zhang & gs 0502 (HNWP) (1b); Zhang & Ren 5572 (PE) (1b), 6082 (PE) (1b), 6322 (PE) ) (1b); Zhang. 7. Н. et al. 0097 (HNWP) (1b), 0414 4291 (HNWP) (1b), 4317 (HNWP) (1b), 4320 (HNW P) (1b); Zhang & Zhou 22092 (PE) (1b), 22155 (PE) (1b), 22675 (PE) (1b), 23742 (PE) (1b): Zhao, J. €. 82- 293 (PE, XJU) (2), ye К (РЕ, XJU) (2), s.n. (PE, XJU) (2); Zhen, S. X. 0351 (PE) (1b); Zhou, H. J. 708 (NWTC) (lo Zhou, L. H. дз (HNWP) (1Ь); Zhu, G. А 5665 (РЕ) (2), 5755 (PE) (la), 6345 (PE) (la). 80006 (NW m. 1b); Zhu, G. L. et al. 6325 (PE) (2). 6386 (PE) (2); Zhu, S. F. 20491 (PE) (1b). PHYLOGENY OF SMELOWSKIA AND RELATED GENERA (BRASSICACEAE) BASED ON NUCLEAR ITS DNA AND CHLOROPLAST trnL INTRON DNA SEQUENCES! Suzanne I. Warwick,” Ihsan Al-Shehbaz.? Connie А. Sauder," David F. Murray,” and Klaus Mummenhoff" ABSTRACT sing sequence data from the ITS region (internal transeribed spacers ITS] а ITS2 of nuclear ribosomal DNA and dhe 2.85 rRNA gene) and ns DN A sequence data from the tral, intron, we examined the evolutionary re ема of and New World Smelowskta species with its putatively re nete genera (Brassicaceae). ITS and ті. sequence dat e obtained from all 13 Sn ielowskia species an id from 13 taxa of the related genera Descurainia, ae. Gorodlriu Hedinia, Polyctentiun, Redowskia, Sinosophiopsis, and Sophiopsis. Results Е maximum parsi- mony analyses s g plus Ermania. Corodko wia, Hedinia, Redowskta, whereas the other clade included Smelowskia holmgrenii | subelade). Descurainia formed a separate cla Gorodkovia, and Redowskia, not a spe C le = Were e xamine xd i In these ge ne ra) genera Ermania. rowed a polyphyletic origin for Smelowskia, with the taxa spli t into two major clades. Smelowskia taxa н тө and Sophiopsis formed one clade (Smelowskia s.l. clade), (Holmgre ni sube lade) an id the genus Polyctenium (Polyctenium le. ITS and trib molecular data, along with a comparison of key morpho- logical traits for each genus, support recognition of a single genus that would include Smelowskia, th e three monotypic as well as the genera Hedinia, Sinosphiopsis, and Sophiopsis (although . The molecular data are consistent v i vith an Asian origin for Smelowskia ‚ followed by diversification in Asia Бай North America, subsequent to migration into arctic ык ын regions of western North America. ITS sequence divergence estimates ranged from O to 4.1% within Smelowskia s. str., and up to 0.15% within the Smelowskia s.l. clade; whereas tral sequence divergence estimates ranged from 0 to 1.85% within Smelowskia s. str., and up to 3.71% within the Smelowskta s.l. clade. indicating pleistocenic speciation within Sme- lowskia as a whole. Key words: Вт Smelowskia, Sophiopsis. trnL.. rassicaceae, Descurainia, Ermania, Gorodkovia, Hedinia, IVS, Polyctenium, Redowskia. Sinosophiopsis, The Brassicaceae are a large family of about 340 genera and over 3350 species distributed through- out the world, primarily in temperate regions (Al- Shehbaz, 1984: Appel & Al-Shehbaz, 2002). It is a well-defined family easily distinguished from the related Cleomaceae by floral and fruit morphology (1.e.. cruciform corolla, tetradynamous stamens, and septate capsular fruits (Hall et al., 2002). classification of the family Tribal is. however, controver- sial, and its phylogeny requires clarification (Koch, 2003; Koch et al., 2003). The characters tradition- ally used at the tribal rank are few and include orientation of the radicle in the embryo, number of rows of seed in each locule, calyx orientation, tri- chome type, fruit compression type (length-width ratio), and features of the nectaries. Several tribal systems have been proposed, and most of the tra- ditional groupings, with the exception of the Bras- siceae and perhaps the Heliophileae, are not mono- phyletic (Koch et al., 2001. 2003: Appel & Al-Shehbaz, 2002). Generic delimitation is also a frequently encountered problem (Al-Shehbaz. 1984; Appel & Al-Shehbaz, 2002), and as many other genera of the family, Smelowskia C. A. Mey. is artificially delimited and lacks unique synapo- morphies (Table 1). All species are perennials with ! We thank Herbert Hurka for providing of the herbaria listed in Table 3. We k "D Ottawa, Ontario КТА 0C6, Mis Canada. wat P.O. Bos 2 9. St. T4 in Canada, 80 uri Botanic г I 71010 ' Agriculture and ; Ottawa, Ontario KIA ч University of / Eastern. С чеш = ш em Y 4 y E p A a S D da gs TER Museum Drive, ^ De po of Botany, Germa ni- Каша ck. DE ANN. MISSOURI Bor. lis, 1 63166-0299, U.S.A. and Oilseed Research ( Fairbanks, Fac ulty of Biology/Chemistry, University of Osnabriick, Barbarastr. а sample of Hedinia tibetica. We are grate ful to the directors and curators thank Victoria Hollowell for editorial advice. e and Agri-Food Canada. Easter m TE and Oilseed Research Centre, Central Experimental Farm, ihsan.al-shehbaz@mobol. org. Centre, Central Experimental Farm, Alaska 99775, U.S.A. ffdfm@uaf.edu. 11, D-49069 Osnabrück. GARD. 91: 99-123. 2004. 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Su gu 5 8 = AE 5 = 3 g variously shaped (linear, oblong, obovoid, ellipsoid, 2 = — T 2 3 or lanceolate) fruits, a prominent midvein on the з Е. S 5 valves, entire stigmas, uniseriately arranged seeds, = O E = L T u 3 "г uM . : E: v5 2 T Б 8 and incumbent cotyledons. The majority of species 2 т c — * = » <= D ы J E | Sle ae c Es СЕБЕ have soft, dendritic and forked hairs, herbaceous, кро Ea .® 2— n = <€ E SE cle s z Z È divided leaves soft in texture, and flattened or an- - - — . кы gled fruits. but S. holmgrenii does not have this 8 E Б combination of characters. ES . = С : А "AM Е v E 5 Smelowskia has been variously delimited (taxo- 8 D = -à А HAE EN a Si» . a T E S nomic treatments vary from 7 to 14 species). with aj = E: = B : : BIS 3 E E = several segregate genera placed in subtribe Des- Als 8 5 Ba. © ——— — curainiinae of the tribe Sisymbrieae (Drury & Rol- _ Ф lins, 1952: Botschantzev. 1968; Velichkin. 1979: 2 y ^" = S Rollins. 1993; Czerepanov, 1995: Mulligan, 2001; 2/> 3 3 = = r ‘ Sy Е 31358 E E 2 = Zhou et al. 2001). As currently circumscribed, бЭ | 7-9. m — 7 2 y y 2% q = no : ПАШ ў ; à . ad жр q E > E Smelowskia includes 13 species, of which 7 (5. a am = oe TA m ae * . oe A americana, S. borealis, S. holmgrenii, S. johnsonii. a — S. media. S. ovalis, S. pyriformis) are North Amer- Š E > 5 ай ican, 5 (S. alba, S. bifurcata, S. calycina, S. ino- elu '* б = ee pinata, S. pectinata) are central and north Asian, =|-= = EZ 3 т = j ens ; "|? 8 = E vt and 1 (S. porsildii) occurs on both continents. wi Ф o z om A = . Names of accepted taxa, synonyms, and excluded g E names in Smelowskia are listed in Table 2. Varieties = v 2 = ‚= of S. calycina previously recognized in North Amer- E © = © — і | \ g «5 4 = mel — E um 3 4 . x Sle та c E DN a ica are treated as distinct species. gi 4 3 E б д ж p р - = »- хе d: «== аар, : Р Р " А А S |= S = > S Schulz (1924, 1936) placed Smelowskia in the — Ф — = „= сч * tribe Sisvmbrieae subtribe Descuraintinae. along z ads with Descurainia Webb & Berth.. Hugueninia E له‎ as E <3 6.3 = Rehb.. desa Cham. & Schltdl., Robeschia = D 5.5 — S E і Fie AE S5 E asia Hochst. ex О. E. Schulz. Sophiopsis O. E. Schulz. rls gE ES 2 = == @ 0 = = © E 2. 3 and Trice о. O. E. Schulz. He divided Sme- lowskia into section Prost tes (S. alba, S. caly- x Ш . „ B . * . H я _ LE cina, S. lineariloba. and S. ovalis) and section Po- E v " T * 2 lyctenium (Greene) O. E. Schulz (S. fremontii). In T = — ك‎ on. $ i А Р 4 Зо = = E 2 — North America. Rollins (1938) recognized Sme- ROI = v = + E д : : Р 3 К м А ЗЕ 3 E: © 3 lowskia (S. calycina and S. ovalis) and Polyctenium Greene (P. fremontii) as closely related genera. E Hultén (1945) placed the monotypic genera Me- = v 2 3 5 lanidion Greene (M. boreale Greene) (Greene, 3 © К: — e Elo «= F E E 1912) and Acroschizocarpus Gombóez (A. kolianus SJE £ b 3 <= Sle 6 E z © Gombócz) (Gombóez. 1940) in Ermania borealis _ (Greene) Hultén. However, this transfer was inva- з 2 | = lid. as the generic name Ermania (Chamisso, 1831) “= a 6 < = = — Р . E EE doses a was not validated until 1956 by Botschantzev (re- 7 E orf = 5 д = © . | | . : | E Sjeo 28 8 88 8 5 zo viewed in Al-Shehbaz. 2001). Drury and Rollins e [835 FsS 28 8 5 B ш и igiiur | 2 KE 3 a" ties 23” (1952) supported the congeneric status of Melani- — = Ф 2 = д — | р . À 3 | dion and Acroschizocarpus and placed them in Sme- ES "T Sa lowskia rather than Ermania Cham. ex Botsch., as . = 2 — Sn 5 — ч S © ＋ © ye Es 38 8 SE S. borealis var. borealis and var. koliana, respec- = 3 = — — DO : Р : Р = £ Z oS 9 & №9 5 tively. Rechinger (1954) reduced his (Rechinger. = — — р = 2278599 - "e : е. D h 2 ә 5 1951) monotypic Chrysanthemopsis (C. koelzii Missouri Botanical Garden Annals of the Dnjnuidoui ni4smoj2uiq nu "< AOJIYSOIO 333¥) n2uini 1d0U1 Di3smojauig naui * [1450104 ^N A (P398) 229071 с Dow DUIPIY &xsdr] (иоѕшоц ү) 217201) `$ DUIIÁ]DI DIYSNO]QUIS DIVUDYISUDA `Ç DAUDYISUDN `Ç UYA DIVUDYISUDA `Ç DUDNANUD DIYSNOJIUÇ тоју © D Gap "e^ Du1oAqDO "$ игуцо әд n10f110n Dds © saplouquiAsis sisdorydog “Ys pay ^g Saplorquidsis с suo fuád piysmojawg , & snusofiád & sur[[oH y Anq H M ed ‘S "ppsaod piysmojaug nppsaod ч nppsaod с nppsaod ‘IPA Du134]D2 с ^ozyunf (suroy X ÁMI] ^H ^W) пррѕлоа - DDD u pismojouig DD: d с DDDjwaM d шуцчоцәд (gung) νd dd nspiompo DIAM ‘dey, 4ojfiaand с sapioXuaDd. Dupuis sapio&uipd Diuuirn] uiun[oq (wey) sapro&aapd "< sippao ртуѕтојәшс̧ sippao "$ sao ‘Ç ,DIS22uo» *sippao “SIBA,, souo[ e N spao + юэ1]оЗиош рппрәр DIYOSUOU рппрә} DAJOFUOU с yosng N (Счлоу) рәуодиош с рәш т1у$то]әш‹; Dipaul `Ç трәш ‘IPA DUIDATDI `Ç uyog (suroy Y &ma( Hd A) отрәш ^s DUDIMIWD DIYSNOJIUS орфо] "$ DUDIMIWD “IRA DUIDATDI *¢ "PAY 22090] ‘Ç DUDIUIWD DIYSNOJINUÇ pgopupauny ^g DUDJ12UID “IBA DUIIATDI `Ç DNN »qogunaum s DUIIAJDI DIYSMOJIUÇ DUloA]DO `Ç nzjooy `Ç DUTIA]DI `Ç yq» Cy u»oy) 12127204 `Ç nppisaod pisnojouiq nppsaod ^q maozjmf с ui nazun с nuosuyol piysnojaug npopaof лел si]D240Q "$ URSIN ^V Y Huosuyol + uumofiusonn Dum] ‘AO "VD mijo fido pur S njpuidoui DIYSMOJIMS njpuidoul с nppuidoui d шоу (шло) DIDUIdOUT ‘S 1тиәл#ш]оц DIYSMOJIUÇ ттиәл#ш]оц & ттиәл#ш]о "< ттиәл#ш]ої N шо 22u242ugow "$ приошә1[ un11212(70 1]uoumaf uın11212(70q uOS]PA, "S 17uoua1f + punssiapyf sisdorydos шу Y UEM Duissiany e umiofipioo Dui241m/] dey v1ofipaoo с 50% ю1узто]әш; “AQ CV `7 DANNI Ç Dul3KpD2 , i- DUIIATDI `Ç DUIDA]DI `Ç DUIDA]DI с әу сү 77) (u2udois) ,n "6 Do1u40fi]po VDIUIDANISA(] ABI) Y D21u10/1]D2 d оиа migunuaog] 'pnais s7npo124Q "$ ¿SO pa “npppsol 81]0ә109 niysmojauig $1]D210Q с si]na10q "є ‘DUDIJOY "s1]D240Q "SIPA,, SUTTJOY y AINA] ^H ^A (2u2212)) #урәлод e D1D2nfiq Diysmojauig nip2anfiq EN D1D24nfiq © Di2anfiq € "Qos1og (“әрә”) n1D24nfiq 'S Di SL отэтуј ^q "zin pijofruajdsp + отии? sisdonjdog чапу олиир ‘¢ DUDIIIWD DIYSNOJIUÇ DUDIUAWD ^q DUDJ12UID “IBA DUIJA]DO "є ‘qPAY (зәрләң Y Josey) рирәмәшт "6 DIDI DIUIPIY рро ртпрәр "ysog (paqog) 2210177 с риа mzunuiog AIM CV су ошар $ DQ]D Diysmojauig 50% 8 5% "< 50% с [2224 (Ied) 290 "$ Juno sir (C661) ^oueda1az7) (6161) чїўчәцәд (8961) лә2иецэѕоЯ (6661) ешон sardadg (2с61) suroy y Anq 102 "DI4smojautq jo 51ӘЧШӘШ SB JUNOIOR sių} ur рәїЧәәәр ӘЛР (et) 32€Jp[oq ut Sad Some SNOLIBA Aq poziusooo1 se Di3smnopouig jo satoedg ЕА 914], Volume 91, Number 1 Warwick et al. 103 1 ا‎ of Smelowskia Rech. f.) to S. koelzit, which was later treated as a synonym of S. calycina by Hedge ( [n a worldwide treatment of Smelowskia. Velich- kin (1979) recognized seven Asian species (5. alba, S. koelzii, S tianschanica) and seven North S. bifurcata, S. calycina, S. jurtzevii, pectinata, and S. American species (S. americana, S. lineariloba. S. lobata, S. media, S. ovalis, S. porsildii. and S. spa- thulatifolia). Rollins (1993) recognized five North American species of Smelowskta. S. borealis (4 va- S. calycina (4 varieties), S. holmgreni &. Mulligan rieties), ovalis (2 varieties), and S. pyriformis. (2001) recently recognized two additional species, S. johnsonii (formerly S. borealis var. jordalii W. H. Drury & Rollins) and S. media (formerly S. calycina var. media W. H. Drury & Rollins). Mul- ligan’s (2001) new combination of S. media is il- Howey er. legitimate because of its earlier use by Velichkin (1979). The purpose of the present study was twofold: first. to test the monophyly of Smelowskia as delim- ited by Drury and Rollins (1952), Botschantzey (1968). Velichkin (1979), Rollins (1993). and Czer- epanov (1995) (Table 2), with emphasis on North American versus Asian taxa: second, to test the re- lationship of Smelowskia with the North American genus Polyctenium and putatively related genera. Relationships to Smelowskia have been suggested for Descurainia Webb & Berthel.. Redowskia. Sophioposis by Schulz (1924. 1936). Ermania by Drury and Rollins (1952), Hedinia Ostenf. by Ve- lichkin (1979). and Sinosophiopsis Al-Shehbaz and Gorodkovia Botsch. & Karav. by Al-Shehbaz (pers. obs.). For comparative morphology of these nine and genera, see Table 1. Sequence comparisons of the internal transcribed. spacers of nuclear ribosomal DNA and the 5.88 rRNA gene (collectively. ITS region) and the tral, intron region of the chloroplast genome were used to assess evolutionary relation- ships and taxonomic treatments of Smelowskta, ITS sequence data are the most widely used nuclear regions in phylogenetic analyses within families (Baldwin et al. ceae include Arabidopsis (DC.) Heynh. (O'Kane et al. 1997: Koch et al., 1999a; Al-Shehbaz & O' Kane, 2002; O Kane & Al-Shehbaz, 2003). Bras- and other genera (Yang et al., 1999), Car- damine L. (Franzke et al.. 1998). Caulanthus S. Watson and Streptanthus Nutt. (Pepper & Norwood, 2001). (Koch et al., 1999b), Crambe „ (Francisco-Ortega et al., 1999), Draba L. (Koch & Al-Shehbaz. 2002), Halimolobus Tausch and re- (Bailey et al., 200: 1999), Pachyphragma (DC.) 1995). Examples in the Brassica- sica L. Cochlearia 1. N Lepidium L. Rchb. lated genera (Bowman et al. and баата M. Král (Mummenhoff et al.. 200la). Sisymbrium L. (Warwick et al.. 2002). (Koch & Mummenhoff, 2001). Vella L. (Crespo et al.. 2000). and Yinshania Ma & Y. Z. Zhao (Koch & Al-Shehbaz. 2000). data also have potential phylogenetic utility for (Sweeney & Price. Thlaspi | 2 The trnlL intron sequence Cardamine L. and Dentaria L. 2000). Cochlearia (Koch et al., 1990b), and Cardaria (Mummenhoff et al.. 2001b). Scop. (Bleeker et al.. 2002). and Sphaerocardamum S. Schauer and related genera (Bailey & Doyle. 1999). Lepidium ). Rorippa MATERIALS AND METHODS PLANT MATERIAL Leaves (100 mg or less) were sampled from her- barium specimens or silica-gel dried samples of field-collected material (Table 3). Where possible, multiple accessions were assessed for each species. Ingroup species were selected based on available material and to represent the morphological and geographical diversity of Smelowskia and/or related B ge nera. ITS sequences (Table 3) were obtaine d from laxa (13 species) of Smelowskia (05 accessions) ad 13 species (23 accessions) from the purported related genera Descurainia (D. californica (A. Gray) O. E. Schulz. D. pinnata (Walter) Britton. D. sophia (I.) Webb). Ermania (E. Botsch.). Gorodkovia (G. jacutica Botsch. & Karav.). Hedinia (H. tibetica (Thomson) Ostenf.). Polycten- ium (P. fremontii (S. Watson) Greene and P. wil- liamsiae Rollins). Redowskia (R. sophtifolia Cham. & Schltdl.). Sinosophiopsis (S. bartholomewti Al- Shehbaz). and Sophioposis (S. annua (Кирг) O. К. Schulz, S. flavissima (Kar. & Kir.) O. E. Schulz. 5. sisymbrioides (Regel & Herder) O. E. Schulz). The tral, intron sequences (Table 3) were obtained for 13 species of Smelowskia (25 accessions) and 13 parryoides (Cham.) species (17 accessions) from the above genera. Five outgroup taxa were initially included in both the ITS and tral appropriate outgroup(s) was difficult, since with the exception of the tribe Brassiceae, all other tribes in analyses. Selection of an the family are not believed to represent natural groups. Representatives of two subtribes of Sisym- brieae (sensu Schulz. 1936) included as Arabidopsis thaliana (I Heynh. were outgroups, 1.€.. (Arabidopsidinae) and Sisymbrium irio L. (Sisym- along with representatives of three other (Arabideae). Brassica rapa brinae), tribes: Arabis alpina L. (Brassiceae), and Capsella bursa-pastoris (L.) Medik. (Lepidieae. Capsellinae). The ITS sequenc- es of Arabidopsis thaliana. Arabis alpina, В. rapa. C. bursa-pastoris, and S. irio and the trn, sequenc- es of the first two were obtained from GenBank (Ta- Annals of the 104 Missouri Botanical Garden = [LLSOEZAV (МҰ) VSN (OVA) 2889 Хоту DUDIJOY “IBA $1]рәлоф "S "6c == LOSOEZAV (МҰ) WSN (WIV) 6Ё uoumg DUDIJOY “IBA $1]D210Q `$ "gc — 99C0£c AV (NV) VSN (VIV) 1217 Артуу DUDIJOY “IBA $1]D210Q "S ‘Lg — 69SOETAV (МУ) VSN (WIV) 997€ sauof y 3224214 $1]D240Q “IBA s1[D240Q "S “OZ = PLSOEZAV (МУ) VSN (WIV) I8£1 3224214 $1]D240Q “IBA s1]D240Q "S "Cc. C£S0£c AV 89C0£c AV (МУ) VSN (OVA) 88-£6 3007) ў w) / $1]|D240Q “IBA 8109109 "S. “HZ 0£S0£C AV POSOEZAV (NV) WSN (WIV) £0c£ puvjoy $1]D240Q “IBA S1]D210Q "S “EZ COCOEZAV (MA) epeuer) (VIV) P601 494104 $1]D240Q “IRA 5109109 "S "Cc 6ZSOETAV ZLSOEZAV (MA) peur?) 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Volume 91, Number 1 2004 Phylogeny of Smelowskia LESOEZAV ZO9OET AV (NV) VSN (VIV) Z106 quv]g X j хтшлојыха чу “ZO — €6€0€C AV BISSHY (JT) u's aazunf 9 vaoynyz “TG nppsiod "c 19 — PCIOETAY (OVC) Ys сур» ә uio]DN OGH әчп[ CZ nppisaod чу “QO CCCOCC AN €c90t€c AV (QVO) #889 Аат nppisaod чу ‘6G m L6€0€c AV (OVC us jm 1 uosuyof осор Nn 11 nppis4od ^s gc — GOSOEGAV (OW) Hog ann nppisaod ^q A TcS0€c AV LLS0€C AV (VIV) €292 427g nppssod чу 96 SESOEGAV POODEZAV (AD) Cory unus pjpuijood g == S6€0tc AV (OVC) OSE mop?) DISOFUO) “IBN SIJDAO "S ‘pC NES LOSOEGAV (VA) ‘VSN (OW) 6E8F souo[ "Is 51[0ао “IBA SIJDAO `$ "CC VESOEZAV POSOEGAYV (99) epeue?) (OVA) 9001 Plod X sso ләри SPNO 164 SMO S. 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UOXR] pue ‘ou UOTSSIDIP ‘Ou UOISS32)E “ON UOISSIIIY yur U3“) yurgues) “panuruo”) '€ AL Annals of the 106 Missouri Botanical Garden OLZOEZAV 8SCLESAV (2002) ‘TP 19 Pme 0141 VUMIAQUIÁSIS ^€6 81 Z9€Z AV LOSTESAV (2002) le 19 J91MIBAN suojspd-psing i “Z6 LIC9OETAV COSTESAV (2002) "Te 19 Pey DADA nisspag “16 O8IPEOAV 6€C€!€14V ep pausriqndun *zequaug-[y pue auey O риро «рү “06 €LSPLX CECE!) 2661) [f 19 3uey O DUDIJDYI sisdopiqniy g Апола) [puru ESCOEZAV GL9OOEZAV uejsryifer (AD 010102 sapio1quiásis ^g gg CCOECAV LI90€ c AV ue]suezey (ЧП) “U's 4040480109 6g eunf ZI punssiany "s 779 cSC0€c AV 0190€£c AV (AMNH) £ZF028 snoukuouy отии? sisdorydos “9g OSSOETAV 6090€c AV (OW) 000€ ҮШ 19 OH 11‹ пэшојоцнтд sisdorydosourg "68 СҮСОЄСЛУ 8090€Z AV (ЯП) gr] aj mijofinjdos piysmopay “PR 6€SS0€c AV S[90€c AV (OW) 61268 шоу y suo amsumpnn d eg 8€C0tc AV TI90€c AV (OIN) 6€ 12214pu() циошәл/ f c9 LT90€£c AV (OVA) £0ZII T? 12 1uya1] 1иошәл/ q 18 — € [90€c AV (OVd) 2088 ismbuos”) njouaf UMIUIIA]O 08 [SCOEZAV LZOVETAV bie (uinueqiou ou) 00-0S-G0$0-66 yang 02112411 Н OL LO90€EZAV Bur) (OW) ££01 I? 12 oH 021991 DUIPIH ‘BL OVEOEZAV S090€c AV eissn] (VIV) ZSE Артуу o $04 nanan у 22 9190€c AV BISSNY (WIV) S Хоту $ soya noimopf м) ‘OL SPSOETAV 9090€c АУ erssny (ЯП) 1/92 1ysunyspAppy DAMIDÍ DIAOIPOL02) ‘C1 TPSOEZAV 9c90tc AV Pissny (VIV) 0211 124104 sapio&umd 7p y ОРСОК АУ CZOOEZAV eıssny (WIV) ZF09 u1qo4dvg Y Je sopr0A110d тири “El LVSOSGAV 6[190€c AV (As) epeueg (OVA) 69c] smuoup piydos (| "c? OPEDEZAV 8 190€Z AV (JS) epeueg (OVC) 722265 [әу pnydos р ‘1L COO CAN (AN) epeue7) (OVA) ZF8ZI uouaqg Y штод ppuuid р 04 == [Cc90€c AV (NO) epeueg (OVA) fess Hum; y 3ooruzay pypuuid є] 69 €TS0€c AV 0c90€c AV 091X3]A (OVA) 9£€27 yoy pipuuid (| °89 STCO€C AV L[90€£cAÀV (AN) "vis'n (OW) £129 4949] v шут DAULSOÍIDI ү 719 COCA 9[90€£c AV (vo) VSN (OW) РРР T"? 1? ]]|onog ponuofi]po DIUIDINISA] “QO CV90£c AV (МУ) VSN (VIV) 019 42104 sruofiád ч ‘c9 = PVOOEZAV (NV) YSA (VIV) £99 49:04 snusofuid чу {9 == 8190£c AV (МУ) YSA (OW) £106 quv]g X 423404 snusofiuád чу ‘g9 quai SLI urduo orde43024) („шпиефәц) uoroo[o-) uoxej pue ou UOISS322P ou uOISS329P ‘ON цоввәдәү yuegua«) yugquo-) "panurmuor) "€ un, Volume 91, Number 1 2004 Warwick e 107 челин К Smelowskia ble 3). Initial analyses showed that if the above five species were included in the outgroup, the ingroup was not monophyletic, as Arabidopsis (DC.) Heynh. and Capsella Medik. formed part of the ingroup. Subsequent analyses included these five taxa. with Brassica, and Sisymbrium assigned to only Arabis. the outgroup. DNA EXTRACTION DNA was extracted from approximately LOO mg or less of dried leaf tissue using Bio 101 FastDNA Kit (Qbiogene Inc., Carlsbad, CA, U.S.A.) following manufacturers. instructions. The final elution. was performed in 80 wl of DES, and then brought to 10 mM Tris (8.0) by the addition of 1/10th volume of 100 mM Tris (pH 8.0). DNA ITS REGION OF NUCLEAR RIBOSOMAL The entire ITS region (including internal tran- scribed spacers ITS! and 1752 of nuclear ribosomal DNA and the 5.85 rRNA gene) was amplifie «das a single unit using primers 151-185 described i O Kane et al. (1997) and I'TS4 described in W lius et al. (1990). The amplifications were performed using Ready-To-Go PCR Beads (Amersham Biosci- ences, Piscataway, NJ. U.S.A.) and 2.0 pl of diluted genomic DNA in a total of 25 pl. were performed on a Thermolyne an ae ther- Amplifications mocycler (Techne Inc Princeton, NJ. Û ) using an initial denaturing step of 95°C b 3 min.; 35 cycles of is 48°C for 35 sec., and 70°C for | min.: and a final extension at 72°C for 10 min. n СК reaction was run on a 1.0% TBE NuSieve (Biowhittaker Molecular Rockland, ME. U.S.A.) agarose gel followed by band excision and purification using the GFN-PCR DNA and Gel Band Purification Kit (Amersham Biosciences). Direct primers (8-88 and ITS4) was performed by for 35 sec., Applications, sequencing using fluorescently labeled Canadian Molecular Research Services (Ottawa, ON, Canada) on a LICOR automated sequencer. Base readings for both forward and reverse se- quences as well as an edited alignment for each sample were verified with associated chromato- grams. The ITS] and ITS2 sequences of accessions 3. 26. 37. 53. and 61 (Table 3) were obtained fol- lowing the protocol given in Mummenhoff et al. (1997 trni. INTRON The trnl. intron was amplified using the e and d primers described Taberlet et al. (1991). The PCR Ready-To-Go PCR Beads (Amersham Biosciences) and 0.2 pl of genomic DNA in a total of 25.0 wl. Amplifications were per- reactions used formed on a Thermolyne amplitron thermocycler Princeton, NJ. U.S.A.) using an initial 39 cycles of (Techne Ine.. 5 step of 94°C for 3 min:; i 45 sec., and 72° for 45 sec.: and a final extension at 72° for 10 min. The CR 1.0% TBE NuSieve (Biowhittaker Molecular Applications) agarose gel followed by purification with the GFX PCR DNA and Gel Band Purification Kit (Amersham Biosci- J for P min.. 50° for reaction was run on a ences). Direct sequencing was carried out on a LICOR automated sequencer using the DYEnamic Direct ith 7-deaza-dGTP (Amer- sham Biosciences) and with fluorescently labeled eyele sequencing kit with primers e and d. Both forward and reverse images were captured and base-called with E-Seq (LiCor. Lincoln, NE. U.S.A.) software. Forward and reverse sequences for individual samples were aligned and edited using AlignIR (Licor). DATA SCORING AND ANALYSIS The sequences were automatically assembled us- ing the software ASSEMBLYLIGN (Kodak. New- haven, CT. U.S.A.) and aligned using clustal in ME- dui (DNA Star, Wl. USA.) and further aligned by eve. For ITS, insertions/deletions (indels), all of which appeared ITS] or ITS2, were treated as missing data in accessions Madison. either — acking the sequence, with the exception of a 16 bp indel in IPS]. The latter indel formed part of a 19 bp region that was excluded from the analysis. This approach retains phylogenetic information from taxa not missing data at the indel. Boundaries of the ITS] and ITS2 regions were determined by comparison with the published sequences of Ara- 1997), Sisymbrium (Francisco-Ortega et al.. 1999), and 2002). It has been previously shown that indels in the tral. intron pro- bidopsis thaliana (O'Kane et al. altissimum L. Sisymbrium irio (Warwick et al.. vide reliable phylogenetic resolution (Mummenhoff et al., 2001b). letions (indels) as missing data, and those indels of 2 bp and greater (10 indels in total) were treated Therefore, we treated insertions/de- as separate characters in the analysis and. scored for presence/absence. Maximum parsimony analyses of the aligned se- quences of ITS and trnL were conducted separately using the computer program PAUP*, version 4.0b2 Swofford, 1999). For the ITS most parsimonious trees were generated using the sequence data, the — heuristic search algorithm, which employed tree bi- 108 Annals of the Missouri Botanical Garden section-reconnection (TBR) branch-swapping. equal weighted characters, with gaps treated as missing data, with 25,000 random additions of the sampled taxa, with 20 trees saved per replicate. Limitations of computer memory required con- straining maximum tree number at 40,000. A sec- ond heuristic search was conducted using the 50% majority-rule tree obtained in the above analysis as the starting tree and the replicate allowed to swap to completion using the TBR option. Statistics re- lating to the amount of homoplasy in the trees, de- scribed in PAUP*, were obtained for each tree and a strict consensus tree was computed for each of the two heuristic analyses. A third heuristic search on the ITS sequence data was conducted where Smelowskia was constrained to monophyly using the constraints function in PAUP*, i.e., 2000 rep- ТВК swap- ping, and a maximum of 20 trees per replicate. Two statistical tests available in PAUP*, the parametric Kishino-Hasegawa and the nonparametric Temple- licates with random addition of taxa. ton. (Wilcoxon signed ranks) and winning-sites tests, were used to evaluate the difference in length between single most parsimonious trees from each of the two analyses. For the trnL sequence data, the most parsimonious trees were generated using the heuristic search algorithm, which employed TBR branch-swapping, equal weighted characters, with gaps 2 bp or less treated as missing data and gaps greater than 2 bp in length treated as separate char- acters (scored presence/absence), with 10,000 ran- dom additions of the sampled taxa, with 1000 trees saved per replicate. In addition, a heuristic search on the trnL sequence data was conducted where Smelowskia was constrained to monophyly using 10,000 rep- TBR swap- ping, and a maximum of LOO trees per rep. ‘he ITS and trnL sequence data were combined the constraints function in PAUP*, i.e.. licates with random addition of taxa. for 47 accessions for which trnL data was available. In order to test whether the data sets were congruent and could be combined, heterogeneity among each data set was assessed using the partition homoge- neity test (Farris et al., 1995) as implemented PAUP. No significant heterogeneity was detected (P = 0.52), indicating that the two data sets could be combined. The most parsimonious trees were gen- erated using the heuristic search algorithm, which employed TBR branch-swapping, with equal weight- ed characters, with gaps treated as missing data for ITS, and for trnL gaps 2 bp or less were treated as missing data and gaps greater than 2 bp in length treated as separate characters (scored presence/ab- sence), with 25,000 random additions of the sampled taxa, with 20 trees saved per replicate. For the ITS, trnL, and combined ITS/trnL se- quence data, 100, 500, and 500 bootstrap (Felsen- stein, 1985) replicates, respectively, were generated in PAUP* using a full heuristic search, with options ACCTRAN, MULTREES = MULPARS, and TBR, and each replicate generated with simple addition sequence of taxa, in order to test the stability of particular nodes in the parsimony analyses. Limi- tations of computer memory required constraining the maximum number of trees to 400 per replicate. Pair-wise distance sequence divergence of ITS and trnL were calculated for each accession pair in PAUP* using the Kimura two parameter model (Ki- mura, 1980) and the pair-wise deletion option for gaps and ambiguous data. RESULTS AND DISCUSSION ITS SEQUENCE DATA A total of 36 taxa (93 accessions) was included in the analysis (Table 3), 5 of which were included in the outgroup. The resulting multiple alignment of the internal transcribed spacer (ITS) region, in- cluding 5.85 gene, was 644 bp long. Sequences are deposited in GenBank (accession numbers in Table 3), and the full alignment is available from the first author. One region of ITS1 (19 bp) was difficult to align and was excluded from the analyses (bases 109 to 127). For the remaining 625 characters, 389 base positions were constant and uninformative, 81 were variable but not parsimony informative, and — 55 were potentially parsimony informative. The ITS] and ITS2 regions exhibited similar amounts of variation, with 129 variable in 260 sites (50%) for ITS] and 99 variable in 201 sites (49%) for ITS2. Only 8 of the 164 sites (5%) in the 5.85 gene showed any variation. The sequence alignment required the introduc- tion of 13 indels in ITS] and 11 indels in ITS Most indels were 1, 2, or 3 bp. However, a 7 bp deletion in ITSI was detected in Brassica rapa, and appears to be the same 10 bp deletion described for its sister species B. oleracea L. Ortega et al. (1999) A 16 bp deletion occurred in the 19 bp ITSI region that was excluded from the by Francisco- analyses and is the same 16 bp deletion reported for Sisymbrium altissimum by Francisco-Ortega et al. (1999) and other Old World Sisymbrium taxa (Warwick et al., 2002) Among the ingroup taxa, the longest ITS region (609 bp, excluding indels) occurred in Smelowskia holmgrenii and the shortest (581 bp, excluding in- dels) in Hedinia tibetica. The G + С content of taxa within the ingroup was similar, with an average val- ue of 55.9%. — Volume 91, Number 1 2004 Warwick et al. 109 Phylogeny of Smelowskia Conspecific accessions of several species of Sme- lowskia. Descurainia, Gorodkovia, and Polyctenium had identical sequences over the entire length of the ITS alignment: these are indicated in parenthe- ses after the species name in Figures | and 2. Iden- tical sequences were also detected for some Sme- lowskia species (Fig. 2). For example. two accessions of S. borealis var. koliana (acc. 28. 29) had identical sequences to 5. media (acc. 4951). ITS sequence divergence (excluding ambiguities) among conspecific accessions was usually small: Smelowskia alba (3 acc., range 0.17-0.34%). S. americana (13 acc.. 0-0.85%). S. borealis var. 0—3.109€). S. borealis var. bo- realis (9 ace.. koliana (Gombócz) W. H. Drury & Rollins (3 ace.. 0— 0.179€). S. calycina (6 acc., 0-3.23%). S. holm- grenii (4 acc.. О-0.67%). S. inopinata (2 acc.. 0%), 0-0.17%). S. media (7 ace., 0— 0-0.84%). S. pyriformis Descurainia e (2 acc., 0%), 5%). D. sophia (2 асс. 0%). Ermanta de j^ 0.33%). kovia jacutica (З ace.. 0—0.509€), Hedinia tibetica (2 ace., 1.39%), and Polyctenium fremontii (3 acc., 0%). The equal weight maximum parsimony analysis, S. Johnsonit (3 acc.. 0.67%). S. porsildii (6 ace., (4 acc., 0%), D. pinnata (3 асс. acc., Tue: = which excluded autapomorphies. yielded 19,507 most parsimonious trees of 417 steps. with Arabi- dopsis thaliana, Arabis alpina, Brassica rapa, Cap- sella bursa-pastoris, and Sisymbrium irto as the out- group taxa. When the 50% majority-rule tree of the above 19,507 trees was used as the starting tree in a subsequent heuristic search and then swapped to completion, a total of 19.739 most parsimonious trees was obtained. Homoplasy in the data set is low as measured by a consistency index (CI) of 0.54 and retention index (RI) of 0.79. The strict consen- sus tree from the two analyses was the same. Figure 39 most parsimonious trees with bootstrap values, and Fig- 1 shows the strict consensus tree of the 19.7 ure 2 shows one of the most parsimonious trees with branch lengths. Four major ingroup clades were ev- ident: Smelowskia s.l. (Smelowskia s. str. and the related genera Ermania, Gorodkovia, Hedinia, Re- dowskia, Sinosophiopsis, and Sophiopsis). Holm- grenii (Smelowskia holmgrenii). Polyctenium, and Descurainia spp. Descurainia was clearly distinct from the rest of the genera included in the ingroup. trni INTRON SEQUENCE DATA A total of 25 Smelowskia accessions and 17 ac- cessions from related genera were included in the analysis (Table 3), along with sequences from the 5 outgroup taxa. The resulting multiple alignment of the Vn intron was 513 bp long. Sequences are deposited in GenBank (accession numbers in Table 3). and the full alignment is available from the first author. A total of 431 base positions were constant and uninformative, 37 were variable but not par- simony informative and 45 were potentially parsi- mony informative. All Smelowskia accessions had a trnl, intron length of 497 bp, with the exception of S. holmgrenii (492 bp. deletion 371—375 bp). The alignment required the introduction of 16 indels to either related genera or outgroup taxa. Five were | bp. | was a 2 bp indel. and 10 were greater than 2 bp. Two 7 bp deletions were observed in Sophiop- sis flavissima (418—424 bp) and in Sinosophiopsis bartholomewii (188-194 bp). А 9 bp (392—100 bp) was observed for the 2 species, thus increasing trnL intron length to 506 — insertion Polyctentum bp. the longest in the analysis. Á unique deletion was observed for S. holmgrenii accessions (371— 375 bp). A unique 5 bp deletion (114—118 bp) and a 59 bp deletion (229-288) were observed for Des- curainia californica and D. pinnata. resulting in the shortest trnL intron lengths in the ingroup (434 bp). The latter deletion (plus 3 additional deletions of 193-228 bp. 289-370 bp. 376—383 bp) was also observed in 3 of the 5 outgroup taxa (Arabis alpina. Brassica rapa. and Sisymbrium irio). Brassica rapa and Sisymbrium irio both had a 10 bp deletion (121—130 bp). The 10 indels greater than 2 characters in the bp were treated as separate analysis. bringing the total number of parsimony informative characters to 55. The G + C content of taxa within the ingroup was similar, with an average value of : б. The following 16 accessions had identical tral. sequences: Smelowskia alba (ace. 1. as listed i Table 3 and Figures). 5. borealis var. borealis (acc. 20. 24). 32). S. S. media (acc. americana (acc. 4. 12). S. calycina (acc. inopinata (асс. 40), S. johnsonii (acc. 42). 48, 49, 51), S. pyriformis (acc. 62). Ermania parryoides (acc. 73. 74). Gorodkovia ja- cutia (acc. 77), Redowskia sophiifolia (acc. 84). The two accessions (43 and 44) of S. johnsonii had iden- tical sequences, as did accessions 71 and 72 of Descurainia sophia. e equal weight maximum parsimony analysis, which excluded autapomorphies. vielded 20 most parsimonious trees of 76 steps. with Arabidopsis thaliana, Arabis alpina, Brassica гара. Capsella bursa-pastoris, and Sisymbrium irio as the outgroup taxa. Homoplasy in the data set is low as measured by a СЇ value of 0.88 and an КІ value of 0.96. Figure 3 shows the strict consensus tree of the 20 most parsimonious trees with bootstrap values, and Figure 4 shows one of the most parsimonious trees 110 Annals of the Missouri Botanical Garden Clades 100 patie alba (1) alba (3) Е 59 Smelowskia borealis var. koliana (27) Smelowskia media (45-4 Smelowskia € bine koliana (28,29) Smelowskia ame а (9 96 e elowskia americana d МА smelowskia americana (1 smelowskia americana 4 8,10,11,14,16) 46) Smelowskia americana 7 с < € Smelowskia porsildii (56) 70 Smelowskia porsildii (61) A Smelowskia porsildii (59 NA Smelowskia porsildii "i 58,60) E € Smelowskia media ) Smelowskia borealis var borealis (2 82 | | — Smelowskia borealis var. borealis (25 Smelowskia borealis var. borealis (2 те! ealis var. borealis دں N N о o 3 2 E о = Ф > о 3 Ф = o < w м d Ф I n = со N © ‹л © о S x А 71 — о o Le is (62-65) Smelowskia borealis var. borealis (21) 90 70 Smelo wskia johnsonii (42) Smelowskia s. l. „ € xmelowskia johnsonii (43,44) RS xmelowskia calycina (30, 34,35) 100 59 Smelowskia bifurcata (1 7) Smelowskia pectinata (55) A 7 ау q |, Li: П íi (22) 7 Frmania narrunidias (73) f J Frmani. irl, (74 ) Smelowskia ovalis var. ovalis (52) Hen s (53) NA Smelowskia „ var. pis (54) Smelowskia inopinata (40,41) Gorodkovia jacutica (76,77) — — 0 8 а (75) Redowskia sophiifolia (84) Hedinia tibetica (78) 100 94 — —— н Hedinia tibetica (79). S Sophiopsis flavissima (87) Sophiopsis annua (86 Sophiopsis sisymbrioides (88) AL Ç < »melowskia holmgrenii (36) 3melowskia holmgrenii (37) Holmgrenii NA 93 ко = A holmgrenii (39 ) P Polyctenium Capsella bursa-pastoris | Arabidopsis/ Arabidopsis thali | Capsella Ag GT , 100 89 Descurainia pinnata (68) Descurainia Descurainia pinnata (69,70) Descurainia sophia (71,72) m Arahis alnina Arabis alpina Brassica rapa Outgroup 3 7 — Figure l. Strict consensus tree of 19,739 most Parsimonious trees based on ITS sequences; tree length = 417 (excluding а ил characters), CI = 0.54, RI = 0.79. Bootstrap ба 's from 100 replicates appear um bran: thes with 9t bootstrap support. Accession number as listed in Table 3 ‘cles name; al Эйн ecific samples listed within parentheses had identical sequences. лө skia s.l. Smelowskia and related genera Ermania, Gorodkovia, Hedinia, Redowskia, Sinosophiopsis, and Sophiopsis|, 5 Holmgrenii, Descurainia) recognized in the analysis and the outgroup taxa are inc dicated to the right. Asian (A) versus North American (NA) distributions are indicated in bold for species in the ingroup. is give n in parenthes ses after ‘he four major clades Volume 91, Number 1 2004 Warwick et Phylogeny B Smelowskia 17 Si perm irio Brassica rapa Arabis alpina 5 changes Figure 2. One of the most vU Ra trees es based o uninformative characters), CI = 0.41. zu four major clades (Smelowskia s 1. 5 Smelowskia calycina (31) A 4, Smelowskia bifurcata (17) melowskia pectinata (55) — — —14 — Sap к ер! Capse z y a calorica (66. 67) Descurainia у Polyctenium. 5 mii. Descurainia) recognized ii . Clades 7 Smelowskia alba (1) 18 > media (45-48 ate realis var. koliana (28,29) ma 9,50,51) i (59 peus loweka porsildii (57. 58,60 1, Smelowskía calycina (30,34,35) Smelowskia s.l. n o = `@ y E^] a melowskia bore: melowskia py е var erai (22) / var. borealis (18, s (74) wskia тора (40,41) Smelowskia er var, nd wskia ova vali Gor Redowskia sophiifol He des иес 78) a tibetica (79) 5 SIS аа (85) ophi 1 Sophio ss p^ ymbriok les (88 Ame я sida reel (36) T ee e hol nii (37) Holmgrenii Smelowskia perii: (38) 1 Smelowskia -— (39) if Polyctenium fremontii (80-82 Polyc eonim fi too (83) bido, рыз thaliana <= Polyctenium ^ | Arabidopsis/ | Capsella 88 © ella burs, a (68 ani a pna (69,70) = Descurainia Outgroup 1 ITS sequences: tree length = 417 (e cluding a The aha of characters supporting each e lade i is indie ‘ated, ı the analysis and the coe taxa are indicated to the right. Asian (A) versus North American (N A) distributions are indicated in bold for species in the ingroup. with branch lengths. Four major ingroup clades were evident: Smelowskia s.l. (Smelowskia S. Str. and the related genera Ermania. Gorodkoria. Hed- inia. Redowskia, Sinosophiopsis. and Sophtopsts). Holmgrenii. Polyctenium. and Descurainia, In con- trast to the ITS data, the trnl. data provided less than 50% bootstrap support (Fig. 3) for the clade containing the Holmerenii. Polyctentum and Ara- Le] e 8 bidopsis/Capsella subclades (Fig. 4 I/ COMBINED SEQUENCE DATA The combined II/, data set included 1143 characters: 830 were constant and uninformative. 112 and The which excluded autapomorphies. yielded 696 most were variable but not parsimony informative, 201 were potentially parsimony informative. equal weight maximum parsimony analysis. parsimonious trees of 461 steps. with Arabidopsis thaliana, Arabis alpina. Brassica rapa, Capsella bursa-pastoris, and Sisymbrium irio as the outgroup taxa. Homoplasy in the combined data set is low as measured by a CI value of 0.60 and an RI value of 0.8 the 896 most parsimonious trees with bootstrap val- . Figure 5 shows the strict consensus tree of ues, and Figure 6 shows one of the most parsimo- nious trees with branch lengths. Analysis of the 112 Annals of the Missouri Botanical Garden Clades p Smelowskia alba (1) | Smelowskia inopinata (40) Smelowskia calycina (32) Smelowskia calycina (35) A Smelowskia pectinata (55) Smelowskia bifurcata (17) es porsildii (56) A Smelowskia porsildii (59) NA Smelowskia borealis var. borealis (20,24) Smelowskia borealis var. borealis (21) Smelowskia s. l. Smelowskia borealis var. borealis (23) i= Smelowskia media (48,49,51) Smelowskia americana (4,12) NA Smelowskia ovalis (52) Smelowskia pyriformis (62) Smelowskia johnsonii (42) 57 Bo cometa johnsonii (43) Smelowskia johnsonii (44) Ermania parryoides (73,74) [— — Redowskia sophiifolia (84) Gorodkovia jacutia (76) Gorodkovia jacutia (77) A Sinosophiopsis bartholomewii (85) { _—————— Hedinia tibetica (79) 100 B annua (86) Sophiopsis sisymbrioides (88) Sophiopsis flavissima (87) Г вене pinnata (68) C D inia californica (66) Descurainia californica (67) Descurainia 95 74 y —Descurainia sophia (71) E . LL Descurainia sophia (72) 99 po holmgrenii (36,38) - | Holmgrenii Smelowskia holmgrenii (39) NA — 100 [^ Polyctenium fremontii (82) Polyctenium Polyctenium williamsiae (83) 83 r— Arabidopsis thaliana Arabidopsis/ Capsella 96 1 Capsella bursa-pastoris 74 [— Sisymbrium irio Brassica rapa Outgroup Arabis alpina Volume 91, Number 1 2004 Warwick et al. 113 19 of Smelowskia combined data set yielded a topology that was near- ly identical to that obtained from the analyses of the ITS data set, with either the same or slightly better bootstrap support. Four major ingroup clades were evident: Smelowskia s.l. (Smelowskia s. str. and the related genera Ermania, Gorodkovia, Hed- inia, Redowskia, (bootstrap support increased from 90 to 97%), Holmgrenii (Smelowskia holmgrenii), Polyctenium, As in the ITS analysis, 5. holmgrenit was very distinct from the Smelowskia — Sinosophiopsis, and Sophiopsis and Descurainia spp. s.l. clade and formed the sister clade to Polycten- ium, although bootstrap support was reduced to 15% compared to 93% for ITS data. RELATIONSHIPS AND CIRCUMSCRIPTION OF SMELOWSKIA l. SMELOWSKIA S. STR. The ITS results indicate that 5melowskia sensu Schulz (1924. 1936) is not a monophyletic genus. Smelowskia species were split into two major clades, with S. holmgrenii (designated the “Holm- erenii” clade) forming a sister group to Polycten- ium. The remaining Smelowskia taxa (Smelowskia s. str.) were included in a single clade (Smelowskia S. I.), along with other genera. The same pattern was apparent for the trnL sequence data. When Sme- lowskia was constrained to monophyly in separate analyses of the ITS and trnL data, using the con- straints function in PAUP* and subjected to the same heuristic search as the unconstrained analy- sis, most parsimonious tree length increased by 43 steps from 417 to 460 steps for the ITS data and increased by 13 steps from 76 to 89 steps for the irnL data. These trees were highly significantly dif- ferent in length (P < 0. 0001) E on both the parametric Kishino-Hasegawa test and the non- parametric Templeton (Wilcoxon signed ranks) and winning-sites test. ле ITS- and trnb-based Smelowskia s. str. clade included S. alba, S. americana, S. bifurcata, S. bo- realis, S. calycina, S. johnsonii, S. inopinata, S. me- dia, and S. pyriformis. With the exclusion of S. holmgrenii, the general ITS- S. ovalis, S. porsildii, r. basi- Rollins based circumscription of Smelowskia s. st cally follows Drury and Rollins (1952), 1993), and Velichkin (1979). The first two ac- counts are essentially identical, but Velichkin rec- ас ognized additional North American species, all of which were treated by the other two authors as va- 2). Velich- kin depended extensively on the degree of devel- rieties or synonyms of 5. calycina (Table opment of nectar glands, abundance of simple versus branched trichomes, and shape of the sep- tum to distinguish most species. However, these characters are so variable even sometimes within a population, that it is highly doubtful if they have any value in the separation of species and were, therefore, justifiably ignored by Drury and Rollins (1952). Asian and North American species did not form separate clades. Based on ITS data, the Asian Sme- lowskia taxa were separated into three well-defined clades, each supported by a 100% bootstrap value (Fig. occurred in 78% of the most parsimonious trees, but had a bootstrap value of less than 50%, and is 1) and a fourth loosely supported clade (which thus not shown in Fig. 1). The first clade included three accessions of S. alba, the second two acces- sions of 5. inopinata, and the third 5. calycina (4 The fourth clade contained two accessions of 5. calycina accessions), S. bifurcata, and S. pectinata. and Ermania parryoides. Smelowskia pectinata and S. bifurcata were sister taxa, separated by only two steps. Morphologically they are also very similar, differing primarily in leaf segment width, which 15 somewhat narrower in 5. pectinata than 5. bifurcata (Velichkin, 1979). The molecular and morphologi- cal data are consistent with the reduction of 5. pec- tinata to synonymy of the earlier-published 5. bi- furcata. With respect to the North American taxa, three species were monophyletic based on ITS data: Sme- lowskia americana (supported by a 96% bootstrap value), S. johnsonii (10% bootstrap value). and S pyriformis (4 accessions with identical sequences " Smelowskia porsildii, which is both North American and Asian, also formed a single clade (70% boot- strap value). The other North American taxa, 5. bo- realis and S. media, were not monophyletic based on ITS data, while relationships of the samples of S. ovalis were not resolved by the ITS data € d 3. e length = 76 (ex cluding parsimony uninformative charac x mee (400 trees per rep) appear above branches with = 500 bo Ta Strict consensus tree of 20 most parsimonious trees oe aa 2 analysis of the trnL intron sequences. 1 M , Cl = RI = P .96. Bootstrap values from 500 El support. Accession number as listed in 3 is given in parentheses after the species name; all conspecific өзө. ч s liste " я тн had identical sequences. The four major clades (Smelowskia s.l., Polyctenium, Holmer analysis i, Descurainia) recognized in the and the outgroup taxa are indicated to the right. Asian (A) versus Noah а (NA) distributions are ibe in bold for species in the ingroup. 114 Annals of the Missouri Botanical Garden ____ Clades | Smelowskia alba (1) | Smelowskia inopinata (40) ¡Smelowskia calycina (32) Smelowskia calycina (35) “таша | Smelowskia pectinata (55) Smelowskia bifurcata (17) 2 Smelowskia porsildii (56) NA Smelowskia porsildii (59) Smelowskia borealis var. borealis (20,24) E | Smelowskia borealis var. borealis (21) 4 smelowskia borealis var. borealis (23) Smelowskia media (48,49,51) ¡Smelowskia americana (4,12) NA Smelowskia ovalis (52) Smelowskia 5.1. Smelowskia pyriformis (62) Smelowskia johnsonii (42) N = Smelowskia johnsonii (43) Smelowskia johnsonii (44) Ermania parryoides (73,74) Redowskia sophiifolia (84) FGorodkovia jacutia (76) Gorodkovia jacutia (77) Sinosophiopsis bartholomewii (85) Hedinia tibetica (79) 8 | Sophiopsis annua (86) Sophiopsis sisymbrioides (88) L2. Sophiopsis flavissima (87) —5— Descurainia sophia (71,72) B Descurainia pinnata (68) 3 + е on Descurainia E N Descurainia californica (66) Descurainia californica (67) — — Smelowskia holmgrenii (36,38) NA Holmgrenii Arabidopsis thaliana Arabidopsis/ Capsella bursa-pastoris _ | Capsella Polyctenium fremontii (82) NA Polyctenium Polyctenium williamsiae (83) 1 Sisymbrium irio Brassica rapa Outgroup — Arabis alpina = Figure 4. One of the most parsimonious trees based on trnL sequence analysis. Tree length = 76 ои parsimony uninformative characters), Cl = 0.88, RI = 0.96. The number of characters supporting each clade indicated. The four major clades (Smelowskia s.l., Polyctenium, Holmgrenii, Descurainia) recognized in the an: as and the outg group taxa are indicated to the right. Asian (A) versus North American (NA) distributions are indic ated in bold for species in the ingroup. Volume 91, Number 1 Warwick et al. 115 2004 1 of Smelowskia Clades Smelowskia alba 0) Smelowskia media (51) NA mel ес calycina (35) Smelowskia calycina (32 Smelowskia borealis var. borealis (20) | Smelowskia johnsonii (42) 87 Smelowskia Johnsoni (43) tien johnsonii (44) N melowskia borealis var. borealis (21) is wskia borealis var. borealis (23) Smelowskia borealis var. borealis (24) пев valis (52) — Smelowskia | 66 = 5 (73) — | 74) Redow skia sophiifolia (84) 99 p— —— Gorodkovia jacutia (75) UH 5 jacutia (77) A [———— Sinosophiopsis bartholomewii (85) 9) LL P tibetica 100 Sophiopsis annua (86) F Sophiopsis кут Аа (88) ا‎ e sima (87) a (6 ә ninn. 523 100 Descurainia californica 100 100 Descurainia californica ja sophia (71 ) Descurainia ph a Smelowskia holmgrenii (36) 100 | Smelowskia holmgrenii (39) Holmgrenii Fi ee Smelowskia holmgrenii (38) N 100 Polyctenium fremontii (82) Polyctenium 1 E (83) = B —— Arabidopsis thaliana Arabidopsis/ Ы———— Capsella a bursa- a Capsella 85 سم‎ a L— ——— Brassic Arabis Bana Polyctenium Outgroup Figure 5. Strict consensus tree of 896 most parsimonious trees generated from analysis of the p ea ITS and tral. intron sequences. Tree length = 461 (excluding parsimony uninformative characters), CI = RI = 0.81. Bootstrap values des 500 replicates (400 trees per rep) appear above branches with > 50% үа support. ecession number as listed in Table 3 is given in parentheses after the species name. The four major clades (Smelowskia sl, Polyctenium. Holmgrenii, Descurainia) recognized in the analysis and the outgroup taxa are indicated to the right. Asian (A) versus North American (NA) distributions are indicated in bold for species in the ingroup. Smelowskia ovalis is morphologically a well-de- two accessions, resulting in a sequence divergence fined species easily separated from the other spe- of 1.34%, ca. 10-fold greater than that observed cies of the genus by having persistent sepals and between accessions of other Smelowskia taxa. Drury ovoid fruits with a distinct style. The accessions and Rollins (1952) and Rollins (1993) recognized corresponding to variety ovalis (ace. 52 and 53) had variety congesta on the basis of inflorescence shape similar ITS sequences, separated by only three (.е., densely congested vs. more lax). silique length steps, whereas variety congesta (acc. 54) was more (3-6 vs. 2-4 mm long), and trichome type (short distinct and separated by seven steps from the other and branched vs. long and simple). In contrast to 116 Annals of the Missouri Botanical Garden Smelowskia alba (1) Smelowskia media (49) Smelowskia media (48) Smelowskia media (51) Smelowskia americana (12) Smelowskia americana (4) A Smelowskia calycina (32) Ermania parryoides (73) Ermania parryoides (74) — Smelowskia borealis var. borealis (20) Smelowskia johnsonii (42) Smelowskia johnsonii (43) Smelowskia johnsonii (44 Hedinia tibetica __Clades |> Smelowskia s.l. ) Smelowskia borealis var. borealis (21) N Smelowskia borealis var. borealis NE Smelowskia ovalis (52) Smelowskia borealis var. н (24) Smelowskia pyriformi Smelowskia 1 (40) Gorodkovia jacutia (75) kovia jacutia (77) ) Sophiopsis a 2 ~ 12 Sisymbrium irio Brassica rapa Arabis alpina 21 22 — 5 changes Figure 6. 461 (excluding ка uninfor clade is indica e four major clades (Smelowskia s.l., analysis and he Mes taxa are indicated to the right. indicated in bold for species in the ingroup. iative characters), CI = the ITS data, which supports the recognition of the two varieties, a critical re-examination of the type material does not support these differences and, therefore, we have proposed the reduction of this variety to synonymy of S. ovalis (Table 2). Based on ITS data, Smelowskia borealis was sep- arated into six clades (Fig. 2). The first contained S. borealis (acc. 27, 28, and 29), all corresponding to variety koliana (Gombócz) W. H. Drury & Rollins —., , Smelowskia holmgrenii (36) ! Smelowskia holmgrenii (39) Smelowskia holmgre 18 Polyctenium fremontii (82) , Polyctenium эмы аы (83) La -— — Arabidopsis thaliana le bursa-pastoris ua (86) ` Sophiopsis pulling ie (88) Sophiopsis flavissim ый 14 scurainia p ta (68) 7 | Descurainia iom (66) escurainia californica (67) Descurainia sophia Descurainia sophia (72) Descurainia Holmgrenii nii (38) ри Arabidopsis/ | Cap sella alla Outgroup One of the most 5 trees based on combined E and trnL sequence analysis. Tree length — 0.60, RI = 0.81. The number of characters supporting each Polyctenium, Holmgrenii, Descurainia) recognized in the Asian (A) versus North American (NA) distributions are and S. media (all accessions). The remaining S. bo- realis accessions (var. borealis) formed loosely sup- ported clades, except for the clade containing ac- cessions 18, 20, and 22, which was supported by a 94% bootstrap value. Smelowskia borealis is high- ly variable, and some of the varieties (i.e., var. dalii) recognized by Drury and Rollins (1952) merit independent status (see S. johnsonii below). Both S. borealis and S. media are diploid (Table 4), but or- Volume 91, Number 1 4 Warwick 117 19 a Smelowskia Table 4. Chromosome numbers for Smelowskia, Ermania, Gorodkovia, and Redowskia taxa reported in the literature. Taxon 2n Country of origin Reference Smelowskia alba 12 Russia Zhukova & Petrovsky (1980) S. alba 12 Russia Zhukova & Petrovsky (1984) S. alba 12 eei : н (1972) S. americana 22 е i AB) Packer ( S. americana 12 U.S O. UT, Drury & px Ж (1952) S. bifurcata 12 s Krogulevich (1976) S. borealis var. borealis! 12 USA. (AK) Murray & Kelso (1997) S. borealis var. jordalii 12 U.S.A. (AK) Dawe & Murray (1979) S. borealis var. koliana 12 U.S.A. (AK) Dawe & Murray (198 1a) S. borealis var. villosa 12 U.S.A. (АК) Drury & Rollins (1952) S. calycina var. calycina? 12 Russia Yurtsev € Zhukova (1972) Zakharyeva & Astanova (1968) S. calycina var. integrifolia 22 USA. (AK) Johnson & Packer (1968) S. calyeina var. integrifolia 12 U.S.A. (АК) Murray & Kelso (1997) S. media 12 U.S.A. (AK) Dawe € Murray (1981b) S. porsildii* 22 U.S.A. (АК) Dawe & Murray (1979) S. porsildii 22 U.S.A. (AK) Hollins (1993) S. porsildii 32 Russia Zhukova & Petrovsky (1984) S. porsildii 18 Russia Zhukova & Petrovsky (1984) S. porsildit 24 Russia Yurtsev & Zhukova (1972) Yurtsev et al. (1975) S. pyriformis 12 U.S.A. (AK) Murray & Kelso (1997) Ermania parryoides 12, 24 Russia Yurtsev & Zhukova (1972) Zhukova & Petrovsky (1984) E. parryoides 12 Russia Zhukova (1980) Berkutendo et al. (1984 E. parryoides 24 Russia Zhukova & Petrovsky (1977. 1980) Gorodkovia jacutica 12 Russia Yurtsev & Zhukova (1972) z. Jacutica 36 Russia Yurtsev & Zhukova (1982) Redowskia sophiifolia 20 Russia Yurtsev & Zhukova (1982) ! Reported as S. borealis. ? Reported as S. calycina * Reported as S. calycina subsp. integrifolia var. porsildii ! Reported as S. calycina subsp. media Drury & Rollins. they differ significantly in the morphology of their caudices. flowers. and fruits. Smelowskia borealis has simple and stout caudices, often branched flow- ering stems. often purple sepals and/or petals, and media has angustiseptate fruits. By contrast. 5. branched and slender caudices, simple flowering stems, white or cream flowers, and terete or angled fruits. Variety koliana, on the other hand, is quite different in fruit morphology (oblong to spatulate vs. obovate) and length (7-18 vs. 3-7(-8) mm long) from variety borealis. and it may well deserve to be recognized as a distinct species. Critical morpho- logical. cytological, and molecular studies аге needed to understand the phylogenetic relation- ships within the S. borealis and S. media complex- es: Smelowskia johnsonii, recently described by Mulligan (2001), would appear to be the same taxon as S. borealis var. jordalii W. H. Drury & Rollins. (Drury & Rollins) Hultén. We have not examined the tvpe of the latter taxon. but the original description is similar to the type collection of S. johnsonii, except the latter. has shorter fruits that are not spatulate. Smeloiwskia johnsonii is morphologically very distinct from all of the North American species and is easily distin- guished by its densely white villous leaves, minute styles (usually less than 0.5 mm long). caducous calyx, and ellipsoid, slightly angustiseptate fruits. The molecular data support the recognition of the varieties of Smelowskia calycina (Drury & Rollins. 1952: Rollins, 1993, Table 2) as separate species as in Velichkin (1979), with S lycina, S. media, and S. porsildii each forming dis- S. americana, 5S. ca- tinct clades. Smelowskia calycina is strictly an Asian taxon, and based on ITS data (Fig. 2) was divided into two clades (ace. 30, ЗІ. 34. 35 and 32, 33). However. no obvious morphological differ- ences were apparent between the herbarium spec- 118 Annals of the Missouri Botanical Garden imens in each clade. The ITS data for accession 35 support the inclusion of S. koelzii in S. calycina by Botschantzev (1968) and Czerepanov (1995). The North American S. media and S. porsildii are dis- tinct cytologically (the former is diploid and the atter is polyploid with various numbers; see Table 4). They also differ in their leaves and fruits. In S. media the cauline leaves are divided and the fruits are widest above the middle, whereas in S. porsildii the cauline leaves are undivided and the fruits are widest at the middle. The remaining North Ameri- can varieties of S. calycina (var. americana (Regel & Herder) W. H. Drury & Rollins and var. integri- (Seeman) Rollins), as treated by Drury and (1952). Rollins (1993), and Mulligan (2001), should be recognized as S. americana, the oldest name available at the specific rank. Velich- kin (1974, 1979) recognized four distinct North American species (5. americana, S. lineariloba, 5. lobata, and S. spathulatifolia; see Table 2) in what is treated here as 5. americana. However, Velich- kin's delimitation of the last species included ma- terial that belongs to S. porsildii. Further studies are needed to elucidate whether or not S. spathu- latifolia merits recognition. 2. SMELOWSKIA S.L. CLADE Representatives of Ermania, Gorodkovia, Hedi- nia, Redowskia, Sinosophiopsis, and Sophiopsis were included within a well-supported clade along with Smelowskia s. str., with bootstrap values of 90 and 1% based on ITS and trnl. sequence data, re- spectively. Except for the three Sophiopsis species, the degree of sequence divergence from Smelowskia taxa for the other five genera fell within the range observed among Smelowskia s. str. Each will be dis- cussed separately below based on ITS and n di- vergence from Smelowskia s. str. Ermania Cham. ex. Botsch. The ITS data fully supported the earlier taxo- nomic transfer by Drury and Rollins (1952) of Er- mania borealis to Smelowskia. These authors fur- ther indicated (1952: 88) that "the type species of Ermania, E. parryoides, also be included in Smelowskia.” might with some propriety Although Polunin (1959) transferred E. parryoides to Smelowskia, Er- mania has been retained by most authors as a sep- arate genus on the basis of having elongated lati- septate siliques, i.e., strongly flattened parallel to the septum. In the analysis of ITS sequence data, H. parryoides formed a sister group to two acces- sions of Smelowskia calycina in 78% of the maxi- mally parsimonious trees (MPT) with a sequence divergence of only 0.17-0.67%. Sequence diver- gence from other Smelowskia taxa was also less than 2.71%, supporting the taxonomic inclusion of E. parryoides in Smelowskia. Data for the trnL in- tron provided similar evidence for its inclusion in Smelowskia, based on sequence divergence of 0— 1.01%. With the exception of accumbent versus in- cumbent cotyledons, all its morphological features can be accommodated in the current circumscrip- tion of Smelowskia (Table 1). Ermania parryoides is 12, 24, as in other species of Smelowskia (Table 4). Therefore, the morphological. cytological, and mo- reported to have chromosome numbers of 2n — lecular data clearly support Polunin's (1959) trans- fer of E. parryoides to Smelowskia. Gorodkovia Botsch. & Karar. Both ITS and trnL sequence data indicate the closeness of Gorodkovia to Smelowskia s. str. Based on ITS data, sequence divergence between Gorod- kovia and Asian Smelowskia taxa ranged from 1.51% to 3.40% and were only 0-1.22% based on rn data. This monotypic genus, described in Bot- schantzev and Karavaev (1959), is distributed in eastern Siberia and the Russian Far East (Czere- panov, 1995). With the exception of having biser- iate instead of uniseriate seed arrangement, all oth- er morphological features of G. jacutica fall within the current delimitation of Smelowskia (Table 1). Similar to the Old World Smelowskia taxa, it has a 12, 36. Morphologi- cal, cytological, and molecular data do not support chromosome number of 2n = the maintenance of Gorodkovia as a genus distinct from Smelowskia. Redowskia Cham. & Schltdl. Both ITS and trnL sequence data indicate the closeness of the monotypic genus Redowskia to Smelowskia s. str. Based on ITS sequence data, sophiifolia was separated from the Asian taxa of Smelowskia by a sequence divergence of only 2.02— 3.76% and 0-1.01% based on trnL data. 1 genus occurs in eastern Siberia (Czere— panov, 1995). Schulz (1924, 1936) placed Redows- kia and Smelowskia in subtribe Descurainiinae. Re- This dowskia differs from Smelowskia in fruit shape, slightly 2-lobed versus entire stigmas, and а versus uniseriate seed arrangement (Table 1). also appears to have a distinct chromosome inch ar of 2n = 20, as compared to 2n = 12, 18, 22, 24, and 32 in species of Smelowskia (Table 4). Despite these morphological and cytological differences, ITS and trnL sequence data do not provide suffi- Volume 91, Number 1 2004 Warwick et al. 119 Phylogeny of Smelowskia cient support for the maintenance of Redowskia as distinct from Smelowskia. Hedinia Ostenf. Both ITS and tral sequence data indicate the closeness of Hedinia tibetica to Smelowskia S. str. On the basis of IT is separated from the Asian Smeloiwskia (5. alba, calycina) by a sequence divergence of only 2.72— 3.95%. Hedinia. contains four species, including one of Hedintopsts S sequence data, Hedinia tibetica a central Asian genus (Table 1). Botsch. & Petrovsky recently reduced to synonymy of Hedinia (Appel & Al-Shehbaz. 2002). Hedinia tibetica was originally described under the illegiti- mate name Hutchinsia (Thomson, 1852) and was later transferred by Lipsky (1904) to Smelowskía. Velichkin (1979) indicated that Smelowskia and Hedinia ave closely related. Except for having brac- teate instead of ebracteate racemes and lacking the — median nectar glands, there are no other characters that clearly separate the two genera (Table 1). In fact. as delimited by Czerepanov (1995), Hedinia has species with bracteate and ebracteate racemes. These two character differences are unreliable for the separation of these two genera, as for most other genera of (Al-Shehbaz. 2000, 2001). Morphologic ‘al and us cular data support Lipsky’s the family transfer о tibetica, the generic type, to Sme- lowskia. Comparative morphological and molecular studies are required for H. altaica Pobed., H. czu- kotica (Botsch. & V. V. Petrovsky) Jurtzev, Korob- kov & Balandin, and H. mongolica (Kom.) Velich- kin. — Sinosophiopsis Al-Shehbaz Within the Smelowskia s.l clade from the ITS analysis (Figs. was the sister taxon to Hedinia tibetica, forming a . 2). Sinosophiopsis bartholomewu loosely supported clade (65% bootstrap value). It was separated from Asian Smelowskia taxa by ca. 13 steps and a divergence of 2.54— 3.77%. Based on tral, data, it is separated by only sequence a steps and a sequence divergence of 0.82— 85%. It is one ol three species recently described : this Chinese genus (Al-Shehbaz, 2000, 2002). Sinosophiopsis differs from Smelowskia by the an- nual growth habit and obscure midvein on the fruit valves and differs from Hedinia in an even greater number of morphological features (Table 1). Further molecular studies on Sinosophiopsis furcata Al- Shehbaz and S. heishuiensis (M. T. Wang) Al-Sheh- baz. are needed to determine the distinctness of Sinosophiopsis from Hedinia and Smelowskia. Sophiopsis О, E. Schulz Although included within the Smelowskia s.l. clade, the three Sophiopsis taxa showed the greatest divergence from Smelowskia s. str.. with a sequence divergence of 4.65-6.45%. Based on both ITS пі, sequence data, this central Asian genus of four (Schulz, 1924: 1952). three of which are included in this paper. was not mono- and species Botschantzev. phyletic. Sophiopsis annua and S. sisymbrioides formed a well-supported clade with a boots strap val- 100% between them, ue of and divergence of only 0.607% separated by 35 steps and sequence divergence of 7.42% from the latter two species based on ITS sequence whereas S. flavissima was data and by 11 steps and sequence divergence of 3.33% differs from the other species of Sophiopsis by hav- based on trnL data. Sophiopsis flavissima ing stems with long, simple or forked trichomes in- stead of dendritic ones, and perhaps merits inde- pendent generic status. Schulz (1924, 1936) placed Sophiopsis and Smelowskia in the subtribe Descu- rainiinae, and Velichkin (1979) indicated that Sme- lowskia was closely related to Sophiopsis. Morpho- logical differences between the two genera аге limited (Table 1). Schulz (1924, 1936) separated Sophiopsis from Smelowskia solely on the basis of seeds that become mucilaginous versus non-muci- laginous, respectively, when wetted, a variable character in many genera of Brassicaceae. Both molecular data sets support the inclusion of So- the genus is not monophyletic and is the most distant phiopis in the Smelowskia s.l. clade. However, in the clade. Further study of the group, including & Vved., is required in order to resolve its closest relatives the fourth species, S. micrantha Botsch. and lack of monophyly. 3. POLYCTENIUM GREENE AND SMELOWSKI/A HOLMGRENII The ITS American Polyctenium (Polyctenium clade) as de- limited by Rollins (1938, 1993), is monophyletic and, together with Smelowskia holmgrenii (Holm- sequence data indicate that the North grenii clade), forms a well-supported clade (93% bootstrap value) that is separated by 32 steps from the Smelowskia s.l. clade. Arabidopsis thaliana and y — > * Capsella bursa-pastoris, initially included in phylogenetic analysis as outgroup species, were sis- ter species to the Holmgrenii and Polyctentum clades in a clade supported by a 91% bootstrap value. Similarly for the trnL data, Polyctentum spp. and S. holmgrenii were included in the same clade (63% bootstrap value). along with A. thaliana and C. bursa-pastoris and separated from the Smelows- 120 Annals of the Missouri Botanical Garden kia s.l. clade by nine steps. Although A. thaliana and C. bursa-pastoris are members of different tribes, several molecular studies (Galloway et al., 1998; Koch et al., 2001) suggest these two taxa to be more closely related than previously thought. The unexpected position of these two genera in our phylogenetic tree within the ingroup as sister to S. holmgrenii and Polyctenium (subtribe Descuraini- inae, tribe Sisymbrieae) underlines the profound problems in tribal classification of the Brassica- ceae. However, this is outside the scope of the cur- rent study. Comprehensive molecular studies along with critical re-evaluation of morphological char- acters are needed to obtain a robust phylogenetic framework that reflects a more natural classification syslem. Rollins (1938) separated Smelowskia from Polyc- tenium by the petiolate, pliable leaves covered with dense whitish tomentum, caudices covered by leaf bases, petals differentiated into blade and claw, and 2 to 10 ovules per ovary. Polyctenium was said to have non-petiolate, wity leaves, sparse, never whit- ish pubescence, caudices free of leaf bases, petals tapering from apex to base, and 12 to 28 ovules per ovary. In his original description of S. holm- grenii, Rollins (1950) expressed the difficulty in as- signing this species to a genus. He placed it in Smelowskia based on fruit morphology (terete, few- seeded). He also indicated that it differed from the other two North American Smelowskia, S. calycina and 5. ovalis, by lacking the white dense tomentum and by having entire, stiff, basal leaves. In addition to their rigid, leathery leaves and almost exclusive- ly simple and forked rigid trichomes, Polyctenium species and S. holmgrenii also differ from Sme- lowskia s.l. by having an obscure instead of prom- inent midvein (Table 1). Smelowskia is essentially an arctic-alpine genus with two areas of concentra- tion, the Cascade and Rocky Mountains of western North America and mountainous areas of central and eastern Asia; Polyctenium inhabits semiarid ar- eas at relatively low elevation in Nevada and ad- jacent California, southeastern Oregon, and south- егп Idaho, whereas S. holmgrenii is a narrow endemic at middle elevations in Nevada (Rollins, 1950, 1993). Velichkin (1979) indicated that Sme- lowskia, including S. holmgrenii, was closely relat- ed to Polyctenium and Capsella, and the molecular data herein do not contradict the association of these genera. Morphological, geographical. and both sets of molecular data, therefore, suggest evi- dence that S. holmgrenii is more closely related to Polyctenium thas to Smelowskia. The ITS and com- bined ITS/trnL data were consistent with the inclu- sion of 5. holmgrenii in Polyctenium; however, the irnL sequence data provided limited support (boot- strap value < 50%) for their placement in a single clade, as shown in Figure 4. The inclusion of Ar- abidopsis and Capsella in the same clade with these taxa (Figs. 1, 2, 4-6), supports the treatment of $, holmgrenii as a separate genus until further studies are completed for this group of related taxa. 4. DESCURAINIA WEBB & BERTHEL. Based on ITS sequence data, Descurainia formed a well-supported clade (100% bootstrap value) that was very distinct from Smelowskia, separated by 45 steps from both the Smelowkia s.l. clade and the Polyctenium clade (Figs. 1, 2). Descurainia was di- vided into two distinct clades for both ITS and trnL data. The difference between the two clades was particularly apparent for the trnL intron with D californica and D. pinnata having two unique in- dels, including a 5 bp (114-118 bp) and a 60 bp deletion (229-288 bp) compared to D. sophia. Des- curainia, a genus of about 40 species, is native to many temperate parts of the world, with important areas of diversity in both North and South America (Rollins, 1993). It is clearly distinct from the rest of the genera included in the ingroup. Morpholog- ically, Descurainia is also very distinct from Sme- lowskia by having glandular papillae, exclusively dendritic trichomes, mostly yellow flowers, and valves without a keel (Table 1). Schulz (1924, 1936) placed both genera in the subtribe Descurainiinae, and the molecular data suggest that Descurainia is very distantly related to the Smelowskia s.l. clade. Molecular data also indicate that D. sophia, the ge- neric type, is distinct from the other two species, and further phylogenetic studies are needed on the genus. In summary, both the ITS and trnL data indicat- ed low levels of genetic differentiation among spe- cies of Smelowskia s. str. (excluding S. holmgrenii), in contrast to a wide geographic distribution (Veli- chkin, 1979: map 169) covering central and north- ern Asia and western North America. The molec- ular-based clade also contained Gorodkovia, Hedinia, Re- and Sophiopsis, and thus supports central Asia as the center of origin of Sme- lowskia s. str., followed by diversification in Asia Smelowskia s.l. Ermania, Asian taxa, dowskia, Sinosophiopsis, and North America, subsequent to migration into arctic-alpine regions of western North America via the former Bering land bridge as was suggested by Rollins (1982). Such patterns were also suggested for various disjunct Brassicaceae genera, among them Stroganowia (K. Mummenhoff, unpublished data) and Lepidium (Mummenhoff et al., 2001b). Volume 91, Number 1 2004 Warwick et al. Phylogeny of Smelowskia 121 ITS sequence divergence estimates, i.e., correct- ed pair-wise distance values based on Kimura-2- parameter model, ranged from 0 to 4.1% within Smelowskia s. str., and up to 6.459€ within the Sme- lowskia s.l. clade; whereas trnL sequence diver- gence estimates ranged from О to 1.85% within Smelowskia s. str., and up to 3.71% within the Sme- lowskia s.l. clade. Recent calculations for several Brassicaceae show that corrected ITS distance val- ues (Kimura-2-parameter model) of 1% correspond to ~ 0.5 to l million years (Koch & Al-Shehbaz. 2002. 2004). within Smelowskia as a indicating pleistocenic speciation whole. 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On the new spec ies from the genus dern skia € ‚ A. Mey. (Cruciferae) in the south-east of the U.S.S.R. “Bot. и. (Moscow & Leningrad) 59: ae 1808. [In Russian. | . 1979, Smelowskia C. ey. (Cruciferae). Crit- ical review and relation to cos genera. Bot. Zhurn. (Moscow : rid grad) 64: 153—171. [In na | Warwick, S ‚ A. Al-Shehbaz, a A. Price & С. Sauder. 2002. шш of Sisymbrium (Brassicaceae) based on ITS sequences of nuclear ribosomal DNA. Canad. J. Bot. 80: 1002-1017. White, T. J., T. Bruns, S. Lee & J. Taylor. 1990. Am fication and direct sequencing of fungal ribosomal RNA li- Volume 91, Number 1 Warwick et al. 123 0 of Smelowskia genes for phylogenetics. Pp. 315— T in M. A. Innis D. H. Gelfand. . Sninsky & hite (editors), Academic Press, | rk. D.-P С & W.-H. Li. of Brassica, Ro- PCR Protocols. Yang, V.- M., K.-N. Lai l 1999. Molecular ой netic Й rippa, Arabidopsis and allied genera based on the inter- nal transcribed spacer region of 188-255 rDNA. Molec. Phylogenet. 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[In Russian. | — A — . 1980. E hne numbe rs and tax- onomy of some species of the . Zhurn. (Mosco ow & Leningrad) 65: i e 50, ln йө | ————, 1984. A po ps study of some spec les pt the family sic Asia. Bot. Zhurn. (Moscow & ое [In Russian. | cience vicis Beijing, and ouis iim hei rs of some . Bot. Zhurn. (Moscow usslan. Press, St. Chromosome ı northern 230-240. PHYLOGENETIC Pedro Torrecilla,?? José-Angel RELATIONSHIPS OF VULPIA López-Rodríguez,? and Pilar Catalán?” AND RELATED GENERA (POEAE, POACEAE) BASED ON ANALYSIS OF ITS AND trnL-F SEQUENCES! ABSTRACT A phylogenetic study of Vulpia, fine-leaved Festuca (Festuca subg. Festuca p. p.), and other related genera of the tribe Poeae (Poaceae) has been conducted using eta ва and combined analyses of DNA sequences from the nuclear ITS region and the chloroplast trnL-F region. All five sections of Vulpia (sects. Vulpia, Monachne, Spirachne, 5 Apalochloa) and three sections and one 5 of fine-leaved Festuca (sects. Eskia, Festuca, Aulaxyper, sect. Exaratae) were included in the analysis. Six minor annual genera that have been considered in the past to be nét either to Vulpia or to these fescues (Psilurus, Ctenopsis, Cutandia, Narduroides, Micropyrum, Wangenheimia) were also analyzed in order to estimate an evolutionary framework for this poorly known group of ephemeral or thin- leaved grasses. Single representatives of other less related groups of Poeae (Festuca subg. Schedonorus and Drymanthele, Lolium, Dactylis, Poa, Aaye Sesleria), Triticeae (Secale), ys Mr hypodieae (Brac hypodium) were also studied. Phylogenetic analyses were performed based on parsimony and Bayesian inference criteria using Brachypodium dis- tachyon as outgroup. “Our results cane ‘ate 1 7 Vulpia and the fine: eme Festuca are, respectively, polyphyletic and paraphyletic assemblages of several taxa. A Жк lei пано group that encompasses the fine-leaved F tuca and Vulpia lineages as well as he: he six ephemeral genera (FEVRE) has ek 1 from both the separate and the Pe aie ITS and trnL-F analyses. Within this FEVRE clade Resa sect. Eskia is shown to be a paraphyletic assemblage basal to the other taxa; the core of the FEVRE clade contains four supported ae representing, respec- tively, the lineages Aulaxyper + Vulpia pro p (diplo id), Festuca /angenheimia, Psilurus /ulpia pro pm (polyploid), and Spirachne + Monachne + Lore re сеа uae ав, constitutes a monospecific linea with apparent but unsupported affinities to some a its potential closer relatives. Representatives of all the Vulpia sections except those belonging to typical section Vulpia j join ina a paraphyletic бе supported assemblage that also includes Cutandia, hom a plicata, and Ctenopsis. Two of the s ae annual genera, Micropyrum and Мий, show distant evolutionary placements in the ITS and trnL-F em . The Festuca subsect. Exaratae taxa belong to a phylogenetic lineage yas from those of the Festuca ovina Du Festuca) and F rubra (sect. Aulaxyper) fescues. Nucleotide substitution rates show significant differences between most perennial and annual lineages; ки ed se- lection is maintained in the slowly diverging ca groups, whereas the rapid adaptive evolutionary processes show by the annual taxa may have pu in an increase ae rate in a. The Mediterranean region is a E as the likely place of diver ergence of these grass lineag words: Сіет opsis, Cutandia, Festuca, FEV RE gı group, ITS, Micropyrum, Narduroides, phylogeny, Psilurus, relative rate test, trnL-F, Vulpia, Wangenheimia. — = € ' The genus Vulpia and other related genera of modern phylogenetic perspective. Single represen- ephemeral grasses, mostly native to the Mediter- tatives of Vulpia have been included, however, in ranean region, have never been investigated from a three recent phylogenetic studies of Festuca, a large 1 We thank Clive A. Stace, Jochen Müller, Robert Soreng, and Jerrold Davis for taxonomic comments and peer revision of the manuscript, Clive A. Stace for 8. чы e of our materials and for providing us seeds of some Vulpia taxa, David Posada and Fernando González-Candelas for their helpful аи оп Bayesian analysis and Relative Rate Tests, Javier Fuertes, се Niet to- Feliner, and Ric ind G. Olmstead for critical review of earlier versions of the manusc ript, Steve Russell for facilitating for us the design of the trnL F ‘fern” forward PNIS Samuel Pyke for helping with the collection of some studied spec па. ii curatorial staff of the herbarium JACA for sending us samples for DNA isolation, and the Conservation Council of the Andalusian Government for facilitating the collection of fresh material and seeds of Festuca taxa at the Sierra три National Park. This work has been supported by a Spanish Ministry of Education and Science grant (BOS2000- 0996 project), by a European LSF grant (NHM, London) to PC, and by a Venezuelan CDCH doctorate fellowship to Departamento de Agricultura, Universidad de Zaragoza, Miguel Servet 177, 50013 Zaragoza, España. Present address: Cátedra de Botánica Sistemática, en Калый de Venezuela, Apdo. Postal 4579, 456323 Мане ‘ay, Estado de Aragua, Venezuela. [See footnote 2 for email address. * Author for c orrespondence. ANN. Missouni Bor. GARD. 91: 124-158. 2004. Volume 91, Number 1 2004 Torrecilla et al. Phylogenetic Relationships of Vulpia genus of perennial species that contains important forage grasses distributed on all five continents. * The pioneer work of Darbyshire and Marwie (1992), based on analysis of chloroplast restriction sites, first established the evolutionary links be- tween Vulpia (V. myuros) and Festuca subg. Festu- ca. Subsequent studies by Charmet et al. (1997) and Torrecilla and Catalán (2002), based on anal- yses of ribosomal DNA sequences, also confirmed the close relationships between Vulpia (V. myuros and V. fasciculata) and the fine-leaved fescues, which include Festuca sects. Festuca, Aulaxyper, and Eskia. These three mentioned works have dem- onstrated that Festuca is a paraphyletic complex and that Vulpia and Lolium fall within different Festuca clades. However, whereas the phylogenetic proximity of Lolium and Festuca subg. Schedonorus (Schedonorus) has been ж EA in depth (Charmet et al., 1997; mation has been obtained eu respect to the re- Саш et al., ‚ very little infor- lationships between Vulpia and Festuca subg. Fes- tuca nad sampling of Festuca representatives was included in the study by Torrecilla and Catalán (2002) in which up to ten different lineages of the genus Festuca were incorporated into the analysis. These authors demonstrated that “Festuca s.l.” (in- cluding Vulpia and Lolium) is monophyletic, the Festuca and the “fine-leaved” Fes- yet split into two diverging groups, so-called “broad-leaved” иса. Within this first group Lolium and Schedon- orus formed a well-supported clade, whereas in the second one Vulpia (V. fasciculata) was associated with Festuca sect. Aulaxyper. The taxonomic treatments of Vulpia, the fine- leaved Festuca, and the six related annual genera have varied over the last two centuries (Table 1). Vulpia has been considered as a part of the genus Festuca by some authors or as an independent ge- nus by others. Linnaeus (1753) first described two of the present species of Vulpia under Festuca, in- cluding the type, Vulpia [= Festuca] myuros, and Vulpia [= Festuca] bromoides. Gmelin (1805) sep- arated Vulpia from Festuca based on its annual abit, long-awned lemma, and very unequal glumes. Hackel (1887) subordinated Vulpia as a subgenus of Festuca. Although some later authors still maintained Vulpia within Festuca, most con- temporary agrostologists have considered Vulpia as a genus on its own (Stace & Cotton, 1980; Clayton & Renvoize, 1986). The circumscription of the ge- nus Vulpia has changed depending on the inclusion or exclusion of different genera, subgenera, or sec- 1981). The Vulpia species differ in their breeding system, num- tions within it (summarized in Stace, ber and size of anthers, spikelet structure, number of fertile/unfertile florets, and shape and size of the lemma callus (Cotton & Stace, 1977; Stace, 1978a). Vulpia sect. Loretia has been considered the most similar group to Festuca as it includes robust an- nuals as well as the three known perennial species of Vulpia and shows strong chasmogamy syndromes (Cotton & Stace, | and Spirachne show intermediate traits between Vulpia sects. Monachne Vulpia sects. Loretia and Vulpia trending toward re- duction in size and increasing self-fertility: these groups encompass both chasmogamous and cleis- togamous species, with sterile distal florets and shorter anthers in the proximal fertile ones. Vulpia sect. Vulpia has been interpreted as the most ad- vanced group. as the majority of its taxa are strong- ly cleistogamous species with single and small-an- thered fertile florets. Section Apalochloa is a quite distinct taxon: its only species (Vulpia unilateralis) also shows a reduced habit and reproductive or- gans, but its flowers are chasmogamous. Apalochloa has been treated as an independent genus (= Nar- durus (Bluff. Nees & Schauer) Rchb.) by several authors (summarized in Stace, 1978a): however, as most of its features are similar to Vulpia it has been 1978a: 198] amended). Cotton and Stace (1976) found the chro- classified within it (Stace, Stace, mosome е number to be fixed within each Vulpia spe- cies je 1) and to show a geographical distri- bution pattern. “Festuca sensu stricto” (subg. Festuca p.p.). as circumscribed by Torrecilla and Catalan (2002), in- cludes at least four distinct groups (as sects. Fes- tuca, Aulaxyper, Eskia, and subsect. Exaratae), though their evolutionary relationships have not been conclusively established yet. These taxa are characterized by their perennial habit and short- awned lemma, traits in contrast to Vulpia, though neither condition is free from homoplasy (Cotton & Stace, 1977). Other taxa previously included in Festuca by Hackel (1882) and Krivotulenko (1960) (sects. Subbulbosae, Scariosae, and Pseudoscario- sae) do not belong to this group as they are mem- bers of the “broad-leaved” Festuca clade (Torrecilla & Catalan, 2002). Festuca sect. Festuca includes the type species Е ovina (within a “Festuca ovina” complex). where- as section Aulaxyper encompasses the red fescues complex). Hackel (1882) classi- fied these groups under one section Ovinae [= Fes- (“Festuca rubra” tuca] though separated into two different series by the possession of exclusively intravaginal shoots (sect. Festuca) versus extravaginal and intravaginal ones (sect. Aulaxyper). Together sections Festuca and Aulaxyper account for most species described Annals of the 126 Botanical Garden issouri M '(peurpropun aie вүәләү Ápiojd пошшоә) (2007) up 9 еоәшој 1000 "е 19 әзиәп] (6861) exuo[q y uspen3iey (9961) JE 19 usnoosury (9261) BRIG у цоцолу, '(Ч8/61 “PBLOL) 21S (0861) 3oquauueqq- padre (SS61) әлеу (7761) 201S y uono?) , xc c piunawquadunga ю1шләиәЗит\ UIE EDIT xp I snanpisq snunpisd u, xe | $әртолпрлю\/ SaPlOANPIDA, SIPIOMPLDN XZ € uma&do421]y umaKdoa421]y umaKdoaoi1]y XZ 9 трирт? DIPUDM() трирт” XZ p sisdouay’) sisdouay) 1938 sisdouay’) XZ I ро]цәою4ү 7129s Dop2oppdy 1938 nop[2oppdy “1998 XZ 9 D11340] “WIS 01]ә107 "1998 De- “Was XZ I auyonaidg 1298 auuonaidg 1598 auyonadg əs xc € JUYIDUO y] 71298 JUYIDUO jy 71228 AUYIDUOJ “1998 XQ K ‘XZ ZI P2 тард 1998 ттд 1998 Dang 129s ттд ттд ттд ттд bin :3qns хр ‘XZ I sido1jmpnosq ‘as sidosppnasg “PƏS DINISA у sappuidipapaju әв sappuidpapajup us хр xX PZ “BO DIYS “1998 DIYSY as AUDA 1298 IDUDA “1998 ADIXIP) [PA soppuizipapa]xz] As хр X eg] “Ro D e "12osqns aDIDIDXY “Woosqns SOJDUIÍDADAJU] “19S Donis, 1998 DJn]sa4 YAS aDIDIDXY “Wasqns „D JOA XOL “XQ 7x9 ‘хр ‘xz 072 Jad AXDINY 71928 4ad&xpgny 71298 D [PA sappuiZpapa]x “198 SIJDULÍDADAIX] лә Xe | ‘XOL xg "x9 "xp "xz 06< home, “oesqns Dome, 'joosqns S2JDULÍDADAJU] "128 SOJDULÍDADAJU] лә DIMISI J YAS Donja “YAS әриппдә] "ioosqns avun) “1998 DU) “1998 bones, "апе Don]sa4 “Bqns bons, “Bqns рәп]$ә J DINISA у D2n]sa 4 D2njsa J Done? гАрто]д XN Apmis в, (2007) (9861) (1861) 2981S (2261) S9^A-1S (2881) [2498 H JO “ON upp) IZIOAUIY (2261) 29818 (2881) PPLH y Bpo], Xy uon X u0107) "R19uo8 рәје[әл ләцо pue ‘dmy ‘Dansa po^eo[-2ug Əy} 103 siogjne juo13jjtp Aq paydope syuəuneaa ЭТШОПОХР |, LH Volume 91, Number 1 2004 Torrecilla et al. Phylogenetic Relationships of Vulpia in the genus. which are widespread in the Northern Hemisphere. Section Festuca presents ploidy levels from diploids to duodecaploids. though the high polyploids are uncommon, most taxa having 2n. = 1х—бх; though showing a similar diversity, is richer in taxa 6x and 8x (10x), and tetraploids and 2x and conversely, section Aulaxyper. with 2n = diploids are very rare (Ainscough et al., 1980). Exaratae was described by Saint-Yves (1922) as a subsection of section Ovinae (= Festuca) based on the strongly invaginated leaf-sheaths shown by 1913). This Hackelian Saint-Yves (1922) to sepa- rate his subsection Exaratae from his typical sub- these plants (Saint-Yves. character was used by $ section Legitimae in Festuca (= sects. Festuca and Torrecilla € Catalán, 2002). The Ex— aratae group fescues are mostly local endemics the mountains. with ploidy levels 2n = 2x and 4x. Aulaxyper: cf. from Mediterranean basin and surrounding Several annual genera of Poeae have been grouped in а Vulpia—-Desmazerta com dex: Vulpia, | I Castellia. Desmazeria. Wangenheimia, Micropyrum, Narduroides. Loliolum, Catapodium, Cutandia. Vulpiella. Sclerochloa (Stace. 1981). Four Ctenopsis. Wangenheimia. Micropyrum. and Cas- g p: Ctenopsis. tellia) were characterized as close allies of Vulpia. two others (Varduroides and Loliolum) as interme- diate between Vulpia and Desmazeria, and the re- mainder as more related to Desmazeria. Desmazer- recently ia, Catapodium, and Cutandia were classified by Soreng and Davis (2000) as belonging to their “Parapholiinae” subtribe. and Sclerochloa as a member of their “Puccinellia complex." ac- cording to their phylogenetic study of Pooideae based on combined analysis of chloroplast restric- tion sites and structural characters. None of those groups were included in their “Loliinae” clade (= Festucineae C. Presl.: sensu Tzvelev. 1982). where Festuca is nested. Ctenopsis is similar to Vulpia in several floral and leaf-blade traits (Paunero. 1963): it has been treated as a section of the latter (Clayton & Renvoize, 1986) though it differs in inflorescence type and hilum length. Crenopsis has been consid- ered to be close to but separated from Vulpia (Cot- ton & Stace, 1977). Cutandia has also been clas- sified as an independent rather different genus from Vulpia (Stace, 1978b). Narduroides. Wangenheimia, and Micropyrum were once included in the hetero- geneous genus Nardurus s. (= Apalochloa p.p.: Stace, 1978a) based on their annual habit and spi- cate to racemose inflorescence. However. these three show enough distinct morphological traits to merit generic status (Stace, 1978a). Clayton and Renvoize (1986) even included Narduroides in the separate tribe Hainardieae Greuter. The latter au- thors considered that Psilurus, a monotypic isolated Mediterranean genus, was close to Vulpia. Vulpia. Ctenopsis, Micropyrum, Narduroides, and Cutandia have been classified as members of the genus Festuca by different authors (summarized in Stace, 1981). been considered close to Vulpia (Stace. 19 Likely these predominantly diploid lineages radi- ated in the Mediterranean area, the region in which Wangenheimia and Psilurus have they are endemic. A second divergence of polyploid Northern-Hemisphere Festuca rubra and F ovina taxa from more ancestral Mediterranean fine-leaved diploids is also considered likely (Kerguélen &. Plonka. 1989; Dubcovsky & Martinez. 1992). In order to test these evolutionary scenarios we studied representatives of the five recognized sec- tions of Vulpia, of four sections and subsections of fine-leaved Festuca, and of the small genera Psi- Cutandia, Narduroides, Micropy- lurus, Ctenopsis. гит. and Wangenheimia (see Table 2). Phylogenetic reconstructions are based on both independent and simultaneous analyses of DNA sequences from the nuclear ribosomal region (ITS1-5.8S-ITS2) and from the chloroplast trnL-F region. eu in- tron—irn-Leu 3’ exon—trnL-F spacer). The ITS re- gion has proved useful for resolving the phylogeny of Festuca and its allies (Charmet et al.. 1997; Саш 2000; Torrecilla & Catalan, 2002) due to concerted evolution (Baldwin et al., 1995) and rate of mutation (Torrecilla & Catalán. et al., moderate 2002). The chloroplast trnL-F region has demon- strated its phylogenetic value in other Poeae groups (Brvsting et al., 2000). the two different genomic data sources is intended The combined analysis of to obtain a consensus phylogeny and to interpret 1 the potential incongruences that might appear i the separate nuclear and chloroplast: phylogenetic reconstructions. a subject of importance to resolve the phylogeny of this group of grasses where retic- ulation has played a major role in speciation. Rates of evolution differ significantly between lineages of fast-to-slow aes ing ar (Bous- 1992: Gaut et al., 1992, 1996; Barra- clough et al... 1996: ا‎ & ш: 2001). This phenomenon is particularly evident for quet et al. the grasses. which include some of the most rapidly evolving within the monocotyledons (Gaut et al.. grass species that propagale sequences 1992). Annual or short-lived perennial rapidly are expected to show higher genomic variability than their re- spective long-lived perennial congeners, suggesting a release from stabilized selection (Саш et al., 1997). Different substitution rates derived from di- verse generation limes or alternative. reproductive levi strategies could. affect the reconstruc tion of phylo- Annals of the 128 Botanical Garden issouri M (ZN) 69 -N u -0d PIS8LPAV LLVSLTAV X01 оГешо(] :єрвләҗ Lag :ереивле) :ureda IY ^N C»en) sisuappaau nonjsa 4 (ZN) 99€10 WIVf C[COLTAV 9LT9LTAV «XQ вәгұ әр ѕәүеиәлу oug :O119A1A :03m] ud, suBuIy-19 prjofiounf bongsa у (Z0) 22 1Ld-WIVf-)d 91882 v 28081 LAV * иеп urs S2[IN391104 :PpeA9N LLII :epeue«) :uredg "wong ^w (JPLH) »2u2q: тәтә 4 (..Хә[Чшоә vagna 4.) 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CH 9!t9!TAV XQ RAOZO'] pupe unde "uueq-a2x1e]y. pIe ]) 1197DU1Y101 тәтә] (Z0) BZ Ld-WIVf-2d ZICBLPAV CG) PSL v AV ög uenf urs so[mzodiog Epey PG peurs шеф SSO SUD NAL DINISO Y bono :so9u94&q song шво LIC9ZTAV EZTPLOLAV (шоо) чете) y PI[[122110], may DomDuastd рәткә 4 A pu S J ¿[9 A9] NONI jENEL “ON UOTSSIIIR учея9чэ^) `рәпиппиоту 2 3[qe Annals of the 130 Botanical Garden issouri M (WOVE) 692€99cf OOT8T LAV 68081 LAV Dra E1023] u :96107) JULIA qur] (18944 7) 9 nos mdma (VOVE) 2666 I£C8ZT AV 06V82 Y AV XG вицирѕиот) гертләс̧ :uredg yury (7]) »20]noru32 vid MA (ЯП) SVO LTOL8PAV L6P82 PAV XG bibi Blof :Sose] aey :үР#пмо,] "uouin(q (eOqsNoYy ag) soməədop mdma (qurq (7]) 070jn21u22 4 =) aanof-jeanog (7]) юурупәтиәЎ “т adÁy) 'sstog (SANO[-[RAN(]) 2172107 “1998 UFD SVD OESEL PAV LAV AG "BISOOIN, YS :snud&?) ÁYosION 29 “sslog 1424Q mdma ( (Чокто 29 'ssiog sd249 4 =) “yoryy sdour 4 2d4) "ssrog. 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АУ XZ O :uredg (1) sean (Puny) spamu vid img (VOVD #6926 COLSELAW — Xp eqaıng әр SPAPN :oduanp[og soning :uredg (є) "uoum(] 57 тард (ZN) 61 Dd FOIgLLAV V608LLAV Xp Joyeusd әр орерәд :ezo3e1e7 :uredg (c) ‘чошт ото vid my (ZN) OL 48-94 АЧТА КЪ 98TBLTAV xp Injoy :ezoze1e7 :ureds (1) oum орото vidma A Tug SLI элә] Ápiojd nos PXE], “ON U0ISSIIIB que gus) `рәпипиоу `g Ә[ЧЕ [, 131 Torrecilla et al. Volume 91, Number 1 2004 Phylogenetic Relationships of Vulpia SOS8LPAV 98882 Fx CESBLEAV S6S8ZTAV VES8L TV 8192814V 1668214 V 20181 LAV 9018 TTL AV LT Peoedy XG SOVBLTAV NG £6v8LVvA VW * LOVELVAY NG КОТАК АЎ NG COVELTAN E 9o0Tt 82 VAV XG 6601821 AV XG 9008 LAV XG COOBT LAY E (IRIS (71) S1714917 vidma =) Ny SUSY :вәәцәлд,] VULI (C00 c) ULJEN у „e, (C %, ee D ¿y 2d4) A0IGOY у цоглорәцәәгу ajouupurtaqq "ап (ZY 21 Ааа TILDA] әр орерәд :PZOSPAIPZ Und (Cu up +” "HV DU1881]]D DINISI] 1 әтә 4 :sdnousin() p pun] niunaquozung (CULL, (Q) Dun og =) qouoo[y nuonsip M ad Aq) Ҷэчэоу DIUNIYUITUDA (WOVE) 86097 f uruldojsy :eosong] cured (CA, y ШИИ А (uno) STLUMOUT y (ZN) LOOLO HIVE epueziy op esayoq :prrpepy сира SOLIpI А SO] E onse әр 89e (ZO) Gele RLYRARA : OBO nde (7П) Poll WIVE ‚р os[epe?) pupe meds TIPUL А? „8 (uenos)) STIL. 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SYGM MEI (лә) pue[2u7] vVOSS8LTAV LOPSOEAV XS (ZOOZ) UWY Y eben "1 auuaıad што] "ү шт1]о7 3U[E") ysy :PUE [FU] €OSBLPAV [cr SOA XG (ZOOZ) че[е1в7) A F[[122110], "png sisuaypad nee (СЧәлцә naopnuipunap Y =) 7] 1011072 с] addy) Yooy ^f ам С^пвәң д) smuouopayog 71228 (Счәлцәс noonumpunap 4 =) 7] 4011042 ᷣꝗ adÁy) "uuo]aq (^neag 4) snuouopayos "3qns Aqua SLI {әлә Арго әэлпос Хеј, "ON UOISS2OOEB Jung uar) "panuguo?) < FEL Volume 91, Number 1 2004 Torrecilla et al. Phylogenetic Relationships of Vulpia 133 genetic relationships via long-branch-attraction and/or increased levels of homoplasy (Bousquet et al.. 1992: Gaut et al., 1992, 1996, 1997). Because Vulpia, the fine-leaved Festuca and the related ephemerals constitute a case study with much di- versity of annual and perennial grass lineages, we have conducted relative rate tests of nucleotide substitutions to clarify the selection of adaptive pat- terns that may have impacted each different lineage (Саш et al., 1997). MATERIALS AND METHODS TAXA SAMPLED AND SEQUENCE DATA Sampling of representatives of Vulpia, the six re- — ated annual genera, and the fine-leaved Festuca was accomplished by collecting fresh leaves and herbarium vouchers. A total of 57 samples corre- sponding to 52 taxa were incorporated into this study. Morphologically variable taxa were se- quenced two (Vulpia muralis, V. fasciculata, V. un- ilateralis. H rubra) or three times (V. ciliata). Eigh- teen of these samples that were previously included in the ITS study of Torrecilla and Catalán (2002) were sequenced for the trnL-F. region and used to complete the sampling of the studied species. One Vulpia myuros ITS sequence deposited by Charmet et al. (1997) in Genbank was incorporated into the study. The full list of taxa with indications of names, authorities, localities, herbarium vouchers, Genbank accession numbers, and ploidy levels is presented in Table 2. The sampling spectrum covered representatives from the five recognized sections of Vulpia (18 sam- ples). the fine-leaved Festuca (24), Micropyrum (2). and one sample each from Ctenopsis, Wangenhet- mia, Psilurus. Narduroides, and Cutandia. Three representatives of the “broad-leaved” fescues, Fes- tuca pratensis (Festuca subg. Schedonorus). F. altis- sima (Festuca subg. Drymanthele). and Lolium per- enne (Lolium), and one representative each from Dactylis. Poa, Puccinellia, Sesleria (Poeae), Secale (Triticeae), and Brachypodium (Brachypodieae) were also studied (Table 2). DNA isolation followed the same procedures stated in Torrecilla and Catalan (2002) implement- ed in some cases with the help of the DNAeasy Plant Mini Kit (QIAGEN). Amplification of the ri- bosomal ITS region (ITS1-5.88-ITS2) and further sequencing were performed as indicated in Torre- cilla and. Catalán (2002) using in both cases the external primers KRC (forward) and ITS4 (reverse). PCR amplification and sequencing of the trnL-F region was conducted using either the external pair primers "c" and “Г” (Taberlet et al.. 1991). or the combined pair "fern" (forward primer: 5'-GGCA- GCCCCCARATTCAGGGRAACC-3). (S. Russell. pers. comm.) and "f" (reverse). 50 pl reactions were prepared with 5 pl 10X buffer, 3 pl MgCl, (50 mM). 4 pl dNTP (10 mM). 1 pl primer 1 (20 uM). I pl primer 2 (20 pM), 34.7 ul ddH,O, 0.5 pl Taq (1.5 units), and 1 pl DNA; the reaction mixtures were subjected to one denaturing cycle of — min. at 94°C, 35 cycles of a 15 sec. denaturing step at 94°C, 30 sec. annealing step at 45°C, and l min. elongation step at 72°С, plus one 7 min. 72°C. The amplified products were purified using the QIAgel extraction kit (QIA- GEN). Purified products were then sequenced with the ABI PRISM Dye Terminator Cycle Sequencing Ready reaction kit (Perkin Elmer) and electropho- termination cycle al resed in an ABI Prism 377 automated sequencer. The direct and reverse sequences of each sample were compared and corrected using the program Chromas. obtaining the respective consensus se- quences. All chloroplast trnL-F samples and most ITS rendered a unique sequence; however, some polymorphic positions were detected in a few ITS samples. Those sites were coded as polymorphisms using the ambiguity IUPAC symbols. A total of 38 new ITS sequences and 57 new trnL-F sequences of Festuca, Vulpia, Wangenheimia, Ctenopsis, Mi- cropyrum, Narduroides, Psilurus, Cutandia, Lolium, Dactylis, Poa, Puccinellia, Sesleria, Secale, and Brachypodium have been deposited in GenBank (Table 2). All but 1 (Festuca indigesta subsp. in- digesta) of the 52 taxa studied have been se- quenced for both the ITS and the trnL-F region. А partial sequence of the ITS region of Festuca ner- adensis (ЇТ®1 spacer, 5.85 gene) was included in the analysis. Each separate set of ITS and trnl-F sequences was first aligned using the algorithm of the program ClustalX (Thompson et al.. 1994) and then adjusted manually using the options of the program Se-Al v. 1.0 alpha 1 (Rambaut, 1996). The boundaries of the ITS region. (ITS1-5.8-ITS2) were determined according to those established by Hsiao et al. (1995) for the Pooideae; those of the trnL-F region (urn intron-trnL 3 exon-trnL-F spacer) were estab- lished according to the chloroplast map of Oryza sativa (Hiratsuka et al.. 1989). Gap indels were coded as binary characters by their presence/ab- sence (0—1 matrix). Only those gaps that were un- ambiguous and potentially informative (Torrecilla & Catalán, 2002) were added to their correspondent sequence matrix and used for phylogenetic analy- sis. 134 Annals of the Missouri Botanical Garden PHYLOGENETIC ESTIMATION, AND SEQUENCE RECONSTRUCTION, RATE DIVERSITY Phylogenetic analyses were based on both Par- simony and Bayesian inference approaches as a means to compare topologies obtained from cladis- tic and nucleotide substitution-model-based meth- ods. The ITS and trnL-F data sets were constructed and subjected to separate analyses. A first set of searches was carried out by both excluding and in- cluding polyploid taxa from the analyses with the intention of clarifying the placement of the poly- ploid lineages with respect to their potential an- cestral diploids by superimposing the polyploids on the diploid-based trees. A second round of analyses was conducted excluding gap characters with the aim of evaluating the phylogenetic signal and influ- ence that ITS and traL-F indels could have in the final trees. Finally, a third seach was accomplished exclusively with the ITS data set excluding the highly variable sequence of Cutandia maritima from the analysis (see Results). All the searches that were conducted exclusively with the diploid data sets resolved the same topologies as those con- ducted with the complete data sets (diploids and polyploids included) for their common taxa (dip- loid) and showed similar levels of support for their respective clades; therefore, only trees obtained from the complete data matrices will be shown here. ITS trees were the same when gap characters were either included or excluded from the analyses: however, an increased lack of resolution was found for some lineages in the trnL-F parsimony-based tree when one informative gap indel was removed from analyses (see Results). Exclusion of Cutandia maritima. from the ITS data set did not alter the resulting topologies or their respective levels of support. Cladistic analyses were performed with Paup* ver. 4.0 beta 10 (Swofford, 2002) following the par- simony criterion. Each data matrix was separately subjected to two heuristic search strategies aimed to find putative isles of equally parsimonious trees. All most parsimonious trees obtained from these separate searches (search #1: closest, TBR, MUL- PARS ON; search #2: 10,000 replicates, TBR, MULPARS OFF, saving no more than 5 trees of score > 10 per replicate) were used to compute the final strict consensus tree. Bootstrap support for the best trees found under the parsimony criterion was estimated through 10,000 1985). The TBR-M (Tree Bisection Reconstruction swapping, MULPARS OFF) strategy of DeBry and Olmstead (2000) was applied for these bootstrap searches in order to get replicates (Felsenstein, random-order-entry of accurate bootstrap values while reducing compu- tational time. Bayesian inference search was later performed for the separate ITS and trnL-F data matrices using Bayes 3.0 beta 3 (Huelsenbeck € Ronquist, 2002). substitutions models was previously conducted on A test of goodness of fit for 56 nucleotide each individual data set using the likelihood ratio test statistic included in the program Model Test ver. 3.06 (Posada & Crandall, 1998) that selects the best substitution model for the given data. The estimated optimal substitution model for each sep- arate data set was then imposed in the subsequent Bayesian analyses. The General-Time-Reversibility model with 6 substitution types was the available option in Mr. Bayes 3.0 for those optimal models (see Results). Separate searches were conducted for each independent data matrix under the GTR mod- (4 gamma rate categories) allowing the program to estimate the likelihood parameters relative to nu- cleotide frequencies, nucleotide substitution rates, and gamma shape (proportion of invariable sites was set to zero concordantly with the best substi- tution models selected by ModelTest for each in- dependent ITS and trnL-F data set). Bayesian analyses were first. performed through 1.000,000 trees generated by the Markov chain Monte Carlo (MCMC) sampling trees every 100 generations; phylogenies sampled from their pos- terior probability distribution were then examined for stationarity in their likelihood values. Sampled points from generations that did not reach a stable equilibrium value (Leaché & Reeder, 2002) were eliminated using the burn-in option of Mr. Ba 3.0. New Bayesian searches of 1,000,000 MC MC generations were conducted for each separate data set; topologies sampled from these new searches were used to construct the respective 50% Major- ity-Rule consensus trees where the percentage of times a clade is recovered is interpreted as an es- timation of robustness. The Incongruence Length Difference (ILD) test of Farris et al. (1994) was calculated to determine if the two data sets (ITS and trnL-F) differ signifi- cantly from random partitions of the same size. The TBR-M approach of DeBry and Olmstead (2000) was applied to compute the significance values of the test replicates. Although significance was achieved from this ILD test (see Results) we decid- ed to unite the two data matrices onto a combined one and to conduct further phylogenetic analyses to explore the resolution obtained from the com- bined analysis and to evaluate topological incon- gruences for lineages differently supported in the and separate and combined trees. The heuristic Volume 91, Number 1 2004 = w сл Torrecilla et Phylogenetic Co ee of Vulpia bootstrap analyses of the combined data matrix were performed using the same strategies described above for the parsimony analysis. Bayesian infer- ence was also conducted on the combined data set following the same procedure as for the indepen- dent data matrices but imposing separate substi- tution models for each partition in the simultaneous analysis using the options provided Mr. Bayes Consensus topologies recovered from the above- mentioned analyses showed differences in branch lengths for several lineages (see Results). In order to investigate the differential rates of evolution that may have been followed by these lineages, we con- ducted relative rate tests of nucleotide substitutions see Results) between different sequences or groups of sequences using RRTree 1.1 (Robinson et a 1998: Robinson & Huchon, 2000). constrains the test between lineages or groups ol m — This program sequences (or between pairs of individual sequenc- es), which are weighted by a given topology. using Robinson & Huchon. — a close outgroup as reference 2000). Rates of substitutions were compared and tested between the main lineages or groups of lin- eages resolved by the Bayesian phylogenetic anal- yses in the separate ITS and trnL-F trees. The test was restricted to the ingroup taxa using Festuca pratensis (Festuca subg. Schedonorus). a member of the close “broad-leaved” Festuca clade. as out- eroup. Comparisons were also extended to pairs of individual sequences within and across lineages with the aim of analyzing the rates of substitutions for taxa that branch lengths with respect to other consectional or close taxa. The Kimura 2-parameter model was chosen to estimate the rates of substitution between the ITS and trnl-F non-coding sequences. Pairwise comparisons between all sequences de- showed substantial differences in tected a few cases. mostly related to the ITS data set. where individual sequences showed increased rates of substitutions with respect to other consec- tional, conspecific, or close lineages (see Results). In order to evaluate if these lineages evolved more rapidly than other lineages or could represent po- tential heterologous samples arbitrarily amplified from direct PCR reactions we analyzed the % G + C content of their ITS] regions. which showed to be the most variable in all the studied taxa. and estimated the minimum-free energies (MFE) of folding of their respective RNA transcripts using the program MFOLD ver. 3.1 (Zuker, 1989; Ma- thews et al., 1999: Zuker et al., 1999). Folding cal- culations were set to the default parameters for RNA sequences (37°С. 5% of thermodynamic op- timatility) and were made at the M. Zuker web site (http://ww w.bioinfo.rpi.edu/applications/mfold/old/ rna/form L.cgi). RESULTS DNA SEQUENCES The ITS from 592 bp (Dactylis glomerata) to 613 bp (Bra- chypodium distachyon), or 599 bp (Vulpia sicula) to 610 bp (Vulpia fontqueriana, V. membranacea. V. unilateralis, Psilurus incurvus) within the ingroup The aligned data matrix consists of 634 po- sequences of the studied taxa ranged laxa. sitions after the introduction of gaps. which ranged from | bp to 4 bp. Three hundred fifty-two out of 634 nucleotide positions are variable (55% of the total): of these 147 are located in the ISI region 3%). 38 in the 5.88 gene region (6%). and 167 in the ITS2 region (269€). Of the variable positions 208 are potentially informative (33% of the total) (94 (15%) in the ITS], 19 (3%) in the 5.85 gene. 95 (15%) in the ITS2). The Vl data matrix ranged from 779 (Puc- cinellia distans) to 908 (Secale cereale) or 819 bp (Festuca rothmaleri) to 848 bp (Psilurus incurvus. Micropyrum tenellum) within the ingroup taxa. The aligned data matrix is made of 1022 positions after the introduction of gaps that varied from 1 bp to 32 bp (a 64 bp deletion in Puccinellia distans). The rate of variability and potential phylogenetic infor- mativeness of the chloroplast trnL-F region is lower than that of the nuclear ITS region for a similar set of taxa. In the trnL-F region 323 out of 1022 po- sitions are variable (32% of the total), and of these 164 (10%) are located in the tral intron, 11 (1%) are located in the tral 3“ exon. and 148 (15%) in the trnL-F spacer. Of 323 total variable sites, 131 are potentially informative (12.5% of the total) (76 (7.3%) in the trnL-F intron, 2 (0.2%) in the iral 3’exon, and 53 (5%) in the trnL-F spacer). PHYLOGENETIC ANALYSIS The ITS Data Set The first heuristic search found one island 213.592 equally parsimonious trees that were 950 steps long. CI = 0.409 excluding uninformative characters, RI = 0.010. dered 6631 equally parsimonious trees of the same The second strategy ren- length and statistics as the previous ones. The strict consensus tree of all these most parsimonious trees is shown in Figure 1. A monophyletic FEstuca sensu stricto + Vulpia + Related Ephemerals (FEVRE) group that encom- passes Festuca sects. Eskia, Festuca, Aulaxyper and subsection Exaratae; Vulpia sects. Vulpia. Monach- 136 Annals of the Missouri Botanical Garden 95 100 50 pescao л N Eyad | 2 . eite brad 100 56 83 = Figure 1. ITS uninformative characters, RI = corresponds to a highly variable sequence of ( Results and Discussion). Festuca rothmaleri Festuca rubra (2) Festuca rubra (1) Festuca juncifolia Festuca rivularis Vulpia fasciculata (1) le ا‎ (2) Cutandia maritima (*) Vulpia AS Vulpia membran Festuca ongiauriculata Festuca ov Festuca ри ИЗ Vulpia ciliata (2) Psilurus incurvus Vulpia unilateralis (1) Vulpia unilateralis (2) Festuca capillifolia Festuca clementei Ctenopsis уп tula Narduroides salzmannii Festuca e Festuca gautieri Festuca eskia Festuca quadriflora lium perenn pate. 1 Festuca altissima Dactylis glomerata Puccinellia distans Poa infirm Se а еа Secale L FL I Ll Br 5 distachyon 0.610). Bootstrap percen tage is indice Schedonorus + Lolium меа c 4 maritima that was both included and е Species numbers correspond to different accessions of the same taxon (see Table 2 — FEVRE Aulaxyper (red fescues) Micropyrum Vulpia p. p. (diploid) Spirachne Monachne Loretia Cutandia Festuca plicata estuca (F. ovina fescues) Psilurus Vulpia p. p. (polyploid) Apalochloa Exaratae Eskia „„ „„ „„ „„ „„ „„ mmm mmm mm mm mm mm mm mm e e e mm mm mm mm Drymanthele tree: strict consensus tree of 213,592 equally parsimonious trees (L = 950, CI = 0.469 exc luding isk m the Pep чи branches. Aster xcluded from analyses ae 2). Volume 91, Number 1 2004 Torrecilla et al. 137 Phylogenetic Relationships of Vulpia ne, Spirachne, Loretia, and Apalochloa; and Psilu- rus, Ctenopsis, Wangenheimia, Cutandia, Micropy- rum, and Narduroides is obtained when Brachypodium distachyon is used to root the trees. The first diverging lineages of the FEVRE group are those corresponding to Festuca sect. Eskia. This taxon forms a non-resolved basal assemblage of five species (see Fig. 1). The core of the FEVRE taxa is also unresolved at the base: five clades collapse with seven single taxa in a polytomy. These termi- nal taxa are those belonging to Festuca subsect. Ё. xaratae (Е borderet, Narduroides, F. clementer, F. capillifolia). pyrenaica, Wangenheimia, and Ctenopsis. Of the five resolved clades one repre- sents the Aulaxyper + Micropyrum + diploid Vul- pia (diploid Vulpia sect. Vulpia) lineage. Micropy- rum is resolved as sister of Festuca sect. Aulaxyper. though this relationship is poorly supported: this group is respectively sister to the diploid Vulpia sect. Vulpia clade. High bootstrap support is shown for the separate lineages diploid Vulpia (bs 100), Micropyrum (bs 99), and the sister taxa Festuca + Ё clade is resolved into two diverging lineages of in- nevadensis rothmalerí (bs 95). The second termixed representatives of Vulpia sects. Spirachne. and Loretia plus Cutandia and Festuca (Vul- pia membranacea + V. fontqueriana)) subclade is well supported, whereas the ((Vulpia brevis + V. sicula). V. geniculata) subclade is mostly robust throughout. the Festuca taxa with relatively high bootstrap sup- port A fourth Psilurus + polyploid Vulpia (polyploid Vulpia sect. Monachne. plicata. The ((Vulpia alopecuros Cutandia). fasciculata), V. A third clade is that of (bs 84) but little internal resolution. Vulpia) clade is also highly supported (bs 99); Psi- lurus is resolved as the closest relative of a clade of Vupia representatives where two sister accessions of V. myuros (bs 83) collapse in a polytomy with two representatives of V. ciliata (bs 81). The fifth clade is that of two representatives of Vulpia uni- lateralis (bs 87) (Vulpia sect. Apalochloa). Model Test found TrN + G to be the best substi- tution model for the ITS data matrix. The Bayesian analysis conducted under the GTR model (nst = 6. ріпу = 0) sampled 8927 trees that reached a stable likelihood value after the burn-in of 1074 trees; the 50% majority rule consensus tree of. all sampled trees is shown in Figure 2. The phylogenetic rela- tionships recovered in this tree are similar to those shown in the parsimony-based tree for the best sup- ported groups (Fig. 1). notably those corresponding to the basal Eskia group, and to the internal clades Aulaxyper + Micropyrum diploid Vulpia, Spi- rachne + Monachne + Loretia + Cutandia + F. plicata, Festuca, and Psilurus + polyploid Vulpia. The Apalochloa clade is nested within a elade of ephemeral taxa (including Wangenheimia, Narduro- ides, and Ctenopsis) in this Bayesian tree. The trnl-F Data Set The first heuristic strategy found one island of 1.094.005 equally parsimonious trees that were 533 steps long, CI = 0.610 excluding uninforma- tive characters, RI = 0.778. rendered 3838 equally parsimonious trees of the The second search same length and statisties as previously. The strict consensus tree of all these most parsimonious trees is shown in Figure 3. The trnL-F strict consensus topology shows a slightly higher resolution for the deep internal branches of the core taxa than that of the ITS strict consensus topology (Fig. 1) though they remain poorly supported. The consensus to- pology was less resolved for representatives of Vul- pia sects. Spirachne, Monachne, and Loretia and related ephemerals and for those of Festuca sub- sect. Exaratae and allies when gap characters were excluded from analysis (results not shown). Four of the five gap characters constitute additional mark- ers for clades already supported by substitution changes: however, a fifth gap shared by represen- atives of Vulpia sects. Spirachne, Monachne. Lor- etia, and Apalochloa plus Cutandia, Festuca pli- cata, Festuca subsect. Exaratae, E pyrenaica and F. clementei is the only character that supports the monophyly of this group in the parsimony-based tree (Fig. 3). Nonetheless. the Bayesian inference. which is solely based on nucleotide sequences (Fig. 4). recovers a similar topology to the parsimony based one. The trnL-F data set also confirms the monophyly of the FEVRE group, which is supported by a high bootstrap value (bs 94) (Fig. 3). Within this clade representatives of Festuca sect. Eskia are paraphy- letic and basal to a core of fine-leaved fescues, Vul- pia, and related annual genera (Fig. 3). In this tral - Е topology, Festuca burnatii and F quadriflora separate earlier from the common ancestor but are unresolved, being followed by the divergence of the remaining Eskia taxa (А eskia, E gautieri, Е ele- gans) that collapse in a polytomy and are sister to the rest. The core of the FEVRE taxa is formed by three lineages that collapse in a further polytomy. The best resolved lineage is the one that encom- passes the "rubra and ovina” fescues and their al- lies (bs 92). Within this group the Aulaxyper diploid Vulpia clade is well supported (bs 97). The clade is sister to the Festuca + Wangenheimia one (bs 92) where the Festuca subclade shows a high bootstrap value (bs 99). Narduroides is resolved as 138 Annals of the Missouri Botanical Garden FEVRE Festuca rothmaleri ensis Mic P tenellum Micropyrum patens Vulpia muralis (1) Vulpia muralis (2) Vulpia bromoides Vulpia fasciculata (1 ulpia fasciculata (2) ulpia brevis «€ Vulpia sicula Vulpia geniculata Vulpia alopecuros Vulpia fontqueriana Me ds membranacea Festuca plic Vulpia unilateralis Vu — Festuca bor — Festuca Dna — Festuca clementei Festuca longiauriculata Cutandia maritima (*) antral mol Apalochloa Festuca (F. ovina fescues) Aulaxyper (red fescues) Micropyrum Vulpia p. p. (diploid) ш $рїгасһпе Мопасһпе Loretia Cutandia Festuca plicata 99 Festuca hystrix Festuca frigida Festuca alpina — - Festus capillifolia 73 Vulpia myuros (1) 99 I Vulpia myuros (2) Psilurus 100 Vulpia ciliata (1) Vulpia p. p. 90 Vulpia ciliata (2) bloi Psilurus incurvus (polyploid) Festuca elegans — Festuca burnatii Festuca eskia Eskia Festuca gautieri SEB BB Bae EEE ERER ow a ann SEB BBB SBSBRB SSBB RBRBRB BBB eeee """"mmuamu 00 olium perenne Festuca pratensis Festuca altissima Dactylis glomerata Puccinellia distans oa infirma Schedonorus + Lolium Drymanthele Sesleria argentea Secale cereale Brachypodium distachyon 0.1 Figure 2. is indicated on the corresponding branche Ast ITS tree: Bayesian 50% e Rule (MR) consensus tree of 8927 trees. Posterior probability percentage erisk corresponds to a highly variable sequence e of C. maritima that was both included and excluded from a "s (see Results and Discussion). Species numbers correspond to different as? Table 2). accessions of the same taxon (see Volume 91, Number 1 2004 Torrecilla et al. Phylogenetic Relationships of Vulpia Un — — = un r2 — Vulpia unilateralis (1) Vulpia unilateralis (2) Vulpia sicula Vulpia alopecuros Cutandia maritima iie borderei a pyrenaica Festuca capillifolia estuca clementei 2 m Vulpia brevis Vulpia fasciculata (1) Vulpia fontquerana bob a membranacea Vulpia geniculata Ctenopsis delicatulá 67 Vulpia bromoides : Festuca rothmaleri Festuca juncifolia Festuca nevadensis Wangenheimia lima Narduroides salzmannii Psilurus incurvus Vulpia ciliata (1) Vulpia ciliata (2) I ا‎ e SEN um perenn Festuca pratensis Festuca altissima actylis glomerata Puccinellia distans Poa infirma Sesleria argent Brachypodium айне оп Secale cereale Figure 3. rs, RI = (0.778). Bootstrap percentage is numbers correspond to different accessions of the same taxon (see indic ö NVI E еее... trn L-F tree: strict consensus tree of 1,694,605 equally parsimonious trees (L uninformative characte ated Table 2) .20£. .0.0000000xX0xX 000 0£. LC CTA FEVRE Aulaxyper (red fescues) Vulpia p. p. (diploid) Festuca (F. ovina fescues) Wangenheimia Spirachne Monachne Loretia Cutandia Festuca plicata Apalochloa Ctenopsis Exaratae Psilurus Vulpia p. p. (polyploid) Micropyrum Eskia Schedonorus + Lolium Drymanthele on the corresponding branches, Spee = 533, Cl = 0.610 exc luding 140 Annals of the Missouri Botanical Garden Leere bci SCR. a 9 Festuca rothmaleri | в = FEVRE Festuca juncifolia в Festuca nevadensis Aulaxyper a : Festuca rivularis (red fescues) s 1 ulpia bromoides a в Vulpia muralis (1) : ا‎ non ie ulpia p. p. и а = жеди Hae в : estuca айша ^ aragonensis Festuca Festuca hystrix (F. ovina Festuca indigesta fescues) stuca frigida ч Festuca glacialis Wangenheimia ; Wangenheimia lima : 80 Narduroides salzmannii . Psilurus incurvus А : : Vulpia ciliata (1) PUE к Vulpia ciliata (2) Vulpia p. p. ] " Vulpia ciliata (3) (polyploid) : : Vulpia myuros (1) : . cropyrum tenellum Micropyrum : Micropyrum patens 2 = 100 Vulpia alopecuros Apalochloa = 55 fb cu Cut ч aridus | = 60 Vulpia bre Spirachne a Vulpia иин (1) Мопасһпе = 1 Vulpia unilateralis (2) | м Vulpia sicula Loretia а 100 r ' a 89 Vulpia fontquerana |Cutandia * : b. Vulpia membranacea WE- - ا‎ ы fasciculata (1) Ctenopsis Vulpia geniculata Festuca Ctenopsis delicatula Е plicata Festuca plicata Festuca clementei Festuca border Festuca pyren Exaratae pyrenaica в Festuca capillifolia : Festuca gautieri a : Festuca eskia И - Festuca elegans Eskia - s Festuca burnatii e А =... Festuca MALA a a « IA A AI 100 Loliu m perenne Festuca pratensis Schedonorus + Lolium Festuca altissima Drymanthele Dactylis glomerata Sesleria argentea Puccinellia distans a infirma Brachypodium distachyon 1 Figure 4. trnL-F tree: Bayesian 50% MR consensus tree of 9890 trees. Posterior probability percentage is indicated on the corresponding brane hes Species numbers correspond to different accessions of the same taxon (see Table 2). Volume 91, Number 1 2004 Torrecilla et al. 141 Phylogenetic Relationships of Vulpia the sister taxon of the former groups but this rela- tionship shows low bootstrap support (bs 56) (Fig. 3). A second lineage of Psilurus + polyploid Vulpia shows relatively high support (bs 85): Vulpia myu- ros is resolved as sister to an unresolved clade of Psilurus and Vulpia ciliata representatives with low bootstrap support (bs 67). A highly robust Micro- pyrum clade includes the sister taxa M. tenellum and M. patens. This clade is resolved as sister to the Psilurus + polyploid Vulpia lineage though this relationship lacks bootstrap support. The third lin- eage includes representatives of Vulpia sects. Spi- rachne. Monachne, Loretia, and Apalochloa. Cutan- dia, Ctenopsis, Festuca plicata. and Festuca sect. F. cle- This group is poorly re- Exaratae (F. borderei, F. capillifolia) plus тете and H. pyrenaica. solved and not strongly supported except for the relationships of the sister taxa Vulpia alopecuros + Cutandia maritima (bs 79). Vulpia unilateralis (Vul- pia sect. Apalochloa) is resolved here as sister to Vulpia sicula though their relationship is not sup- ported (Fig. 3). The goodness of fit test of ModelTest indicated K8luf + G as the best substitution model for the trnL-F The ducted under the GTR model (nst = 6. рту = 0) data matrix. Bayesian analysis con- sampled 9890 trees that reached a stable likelihood value after the burn-in of 111 trees: the 50% ma- M? jority-rule consensus tree of all sampled. trees i shown in Figure 4. The phylogenetic relationships recovered by this analysis are congruent with those based in the parsimony approach, as manifested in the two topologies depicted in Figures 3 and 4. All eroups except the unresolved Festuca sect. Exara- tae taxa plus E clementei and К pyrenaica show congruence in both parsimony and Bayesian trees. The Combined ITSArnL-F Data Set Parsimony strict Consensus topologies and Bayesian 50% majority-rule consensus trees ob- tained from the two separate data sets. ITS and trnL-F.. were coincident for several groups. espe- cially for the best supported lineages Aulaxyper + diploid Vulpia. Festuca. Psilurus + polyploid Vul- pia. and Spirachne + Monachne + Loretia + Cu- tandia + F. plicata and for the basal position of 1—1). that both the nuclear and the chloroplast genomes section Eskia (Figs. These results indicate recover a similar phylogenetic signal for most of the studied taxa. Incongruences between the ITS and trnL-F trees refer to the distinct placement of Vul- pia sect. Apalochloa, as well as Wangenheimia, Mi- cropyrum, and Narduroides. To clarify the evolutionary relationships of those eroups and to improve the resolution of the final tree we decided to unite both data matrices into a combined one and conduct further simultaneous phylogenetic analysis on the enlarged data set. A combined ITS/trnL-F samples that were sequenced for the two molecules: the ILD test of Farris et al. (1994) carried out on this ITS/trnL-F data matrix indicates that the two data sets are significantly different from two random їл data matrix was made of 5: partitions of the same size as the original data ma- trices (1000 replicates; Р = 0.001). about the validity that heterogeneity detected. by Arguments the test between sequence data matrices may have in terms of their combinability have been various (Davis et al., 1998; Johnson & Soltis. 1998: Yoder et al., 2001: Barker & Lutzoni, 2002). Davis et al. (1998). Yoder et al. (2001). and Barker and Lutzoni (2002) considered the ILD test an inappropriate method to test for the combination of data partitions for simultaneous analyses: the latter authors further demonstrated the poor predicting value shown by the ILD test for data set combinability intended to increase phylogenetic accuracy even at critical val- ues as low as P = 0.001 The first heuristic search found one island of 577 equally parsimonious trees that were 1488 steps long. CI = 0.503 excluding uninformative charac- ters. RI = 0.644. The second strategy rendered 686 equally parsimonious trees of the same length and statistics that enlarged the spectrum of equally par- simonious trees. The strict consensus tree of all these most parsimonious trees is shown in Figure 5. The strict consensus [TS/trnL-F tree depicted in Figure 5 is resolved for the main FEVRE groups: this FEVRE clade shows strong bootstrap support (bs 100) and divergence of the basal section Eskia taxa (Fig. 5) resolution is obtained here for the and for the branching of some internal core FEV RE lineages. The two representatives of Vulpia unila- teralis join in a well supported monospecific Apal- ochloa clade (bs 99) that is resolved as sister to Ctenopsis and then to the Spirachne + Monachne + Loretia + Cutandia + F. plicata clade though The Mi- cropyrum taxa also join in a congeneric clade that none of these relationships is supported. shows strong bootstrap support (bs 100). The com- bined ITS/trnL-F tree shows only a partial resolu- tion for the relationships between the core FEVRE groups. The best resolved lineage is that of the Au- laxyper + diploid Vulpia and the Festuca + Wan- genheimia groups, which link together in a clade that is sister to Narduroides; however. these rela- tionships lack bootstrap support. Internal resolution is absent for the remaining lineages. except for a clade that shows a paraphyletic basal divergence of 142 Annals of the Missouri Botanical Garden 12 7 772 22 — —— 2 2 K 2 „ „„ „„ „„ „„ „% „„ % nnn MMB By, a B z Festuca rothmaleri (8x) — FEVRE : 1 Festuca nevadensis (10x) м : Festuca juncifolia (8x) Aul в : Festuca rubra (2) (6x) ee " . Festuca rubra (1) (6x) (red fescues) s : Festuca iberica (6x) : : Festuca rivularis (2x) Vulpia p. p. Р a Vulpia muralis (1) (2x) (diploid) . : Vulpia muralis (2) (2x) : . Vulpia bromoides (2x) "x a : Festuca hy (2x) e = : Festuca longiauriculata (2x) estuca : в Festuca aragonensis (4x) ; a . т ER тга (2х) (Е. ovina fescues) . " Festuca alpina (2x) Wanvenheimi a в Ll genheimia в . 76 Festuca glacialis (2x) . : Festuca frigida (2x) : . Wangenheimia lima 2x) — в : Narduroides salzmannii (2x) : x Vulpia alopecuros (2x) — = s Cutandia maritima (2x) Apalochloa a я Vulpia fontqueriana (2x) Spirachne : = Vulpia membran (2x) : Ы Vul vis (2x) Monachne в 9 Vulpia fasciculata (1) 4x) ; в : Vulpi la Go l Loretia “ ы Vulpia geniculata (2x Cutandia : в Festuca plicata 2x) . в Vulpia unilateralis (1) (2х) Ctenopsis a : Vulpia unilateralis (2) — (2x) | : a Ctenopsis delicatula (2x) — Festuca plicata . м Festuca borderei (2x) = ч в Festuca pyrenaica 4x = А Festuca capillifolia a І Exaratae : = Festuca clementei (2x) — : a Vulpia myuros (1) (6x) — Psilurus = в Vulpia ciliata (1) (43) Vulpia p. p. в E Vulpia ciliata (2) (4x) me . е Psilurus incurvus (4x) — fporplaun : ы Micropyrum tenellum (2х) : : н Місгоругит patens (2х) — Micropyrum н = Festuca elegans 2x) z : Festuca burnatii (2x) = . Festuca gautieri cR Eskia s : Festuca eskia е 2 Sea BEeBBBB BSB Peace Яра ог паш (x) "uuu EIE и пашшшшшшшшпшппшпшпшшпвшпшыш Es 90 со, Schedonorus + Lolium 52 Festuca altissima — Drymanthele 100 Dactylis glomerata Sesleria argente Puccinellia distans Poa infirm Secale cereale E Brachypodium distachyon Figure 5. Combined ITS/trnL-F: strict consensus tree of 686 equally parsimonious trees (L = 1488, CI = 0.503 excluding uninformative characters, RI = Species number ).644). Bootstrap percentage is indica tree; black bars correspond to polyploid lineages ted on the corre аралаш branches. 's correspond to different accessions of the same taxon (see Table = Ploidy levels are mapped into the Volume 91, Number 1 2004 Torrecilla et al. 143 Phylogenetic Relationships of Vulpia the Festuca subsect. Exaratae taxa plus E pyren- aica and E clementei from the topology for the core group of Vulpia taxa and their allies (Fig. 5). The Bayesian simultaneous analysis conducted under the GTR model (nst = 6, pinv = 0) sampled 9643 trees that reached a stable likelihood value after the burn-in of 358 trees; the 50% majority rule consensus tree of all sampled trees is shown in Figure 6. Relationships recovered by this anal- ysis are mostly congruent with those based in the parsimony approach: however, a polytomic assem- blage of Festuca subsect. Exaratae taxa plus Е pyr- enaica and E clementei taxa are resolved here as the the FEVRE whereas the Psilurus + polyploid Vulpia clade and basal lineages of core group. the Micropyrum clade collapse in a polytomy with the “Festuca rubra and ovina” group plus allies. Rates of Substitutions and Sequence Diversity between FEVRE Lineages Values and levels of significance of the relative- rate tests calculated between the main FEVRE lin- enges for the separate ITS and trnL-F sequences are shown in Table 3. Rates of substitutions vary the ITS reflecting the less constrained nature of the тоге for sequences than for the trnL-F ones, ribosomal nuclear region compared to the con- served one of the chloroplast genome. Overall. pe- rennial lineages show significant lower rates of sub- stitution than the annual ones for both ITS and irnL-F sequences; however, most of these low rates are contributed to by representatives of the Eskia group. which show significant differences to almost all other FEVRE lineages for both genomic regions. When the Eskia taxa are removed from the test. the core FEVRE perennial lineages evolve, in general terms, at a slower pace than the core FEVRE an- nual ones but the differences are not significant. Pairwise comparisons between individual sequenc- es show Festuca elegans to be the fastest evolving lineage within the otherwise slow evolving assem- blage of Eskia taxa (results not shown). The core FEVRE perennial groups. also. show marked differences in their respective substitution rates; the Exaratae group evolves slower, followed by the intermediate rate of the Festuca lineage and the more rapidly evolving rates of the Aulaxyper lineage (Table 3). Significant differences in substi- tution rates between the Exaratae group and the ITS and the ul se- quences though they are restricted only to the ITS annuals occur for both the set in several Festuca to ephemeral-groups com- ^ parisons. The Aulaxyper lineage shows higher sub- stitution rates than those of the other perennial firma, 50.5%). groups and evolves only slightly slower than most of the annual lineages except for the Loretia assem- blage ITS sequences. This group evolves faster but not significantly more so than the red fescues if Cutandia maritima is excluded from the analysis (Table 3) (see Discussion). Pairwise comparisons between individual sequences found F rothmalert to be a fast-evolving lineage within the Aulaxyper group with rates significantly higher than those of other consectional taxa like E juncifolia and F ri- vularis in both ITS and trnL-F sequences. Similarly. F. frigida also shows significant higher substitution rates than those of the remaining cosectional Fes- tuca taxa for the ITS sequences and than those of F. alpina and E. clementel for the trnL-F sequences (results not shown). Within the group of FEVRE annuals the Loretta assemblage and Ctenopsis delicatula are the most rapidly evolving lineages with respect to the chlo- roplast V sequences, being significantly faster than the Apalochloa, Micropyrum, and Wangenhet- mia lineages and than the Apalochloa, Micropyrum, and Psilurus + polyploid Vulpia groups respectively (Table cantly faster than the diploid Vulpia and the Nar- The Loretia group also evolves signifi- duroides lineages for the ITS sequences if Cutandia maritima is included in the group but shows no sig- nificant differences to those lineages if this taxon is (Table 3) (see Dis- cussion). Analvsis of pairwise comparisons between excluded. from the calculations individual ITS sequences pointed to Cutandia mar- imd as the fastest-evolving taxon of the FEVRE clade for this genomic region: differences lo rates shown by other taxa were significant in all cases except for the Festuca rothmalert, Vulpia fontqueri- ana, and И membranacea ones (results not shown). The % G + C content and the free energy « folding of the ITS] region showed some differences among the studied taxa. The average percentage of guanine and cytosine residues was 00.5% for the ingroup taxa; however, slightly lower values were observed for the annual taxa (from 50.8% in Vulpia membranacea to 58.7% in V. alopecuros) than in the perennial ones (from 60.4% in Festuca rivularis to Cutandia maritima (54%), this percentage is similar to that of other D de n V. quadriflora). — showed the lowest content of G + C though variable ingroup and outgroup annual species (1.e.. Poa in- The variable perennial taxon Fes- tuca rothmaleri had a percentage of 59%, close to the perennial outgroups F pratensis and Lolium perenne (58%). Average Minimum Free Energy (МЕЕ) of folding of the ISI RNA transcripts was —87.60 kcal/mol (sd = + 6.96 keal/mol) for the ingroup taxa. Folding energies also showed slight 144 Annals of the Missouri Botanical Garden AAA „„ „ „ „ „ AAA AAA DBS 8 и FEVRE 51 Fes 0 um { Ашахурег (гед fescues) Vulpia р. р. (diploid) Festuca (F. ovina fescues) Wangenheimia Psilurus Vulpia p. p. (polyploid) ulp Psilurus incurvus TL Micropyrum tenellum Micropyrum patens ] Micropyrum 00 — Vulpia alopecuros utandia maritima Vulpia fontqueriana Spirachne Vulpia membranacea Apalochloa Vulpia brevis Monachne Vulpia deii (1) Loretia Vulpia | Vulpia MEN Cutandia tuca plicata Ctenopsis Festuca Ctenopsis delicatula SJ plicata — Festuca borderei — Festuca pyrenaica 96 — Festuca capillifolia Exaratae [- Festuca gautieri гү eskia Eskia a burnatii MT 2Q0¿»n£0.%.0X000.L..LAÚLRwf E .s0 0000205000005» б у idi Schedonorus + Lolium aaa altissima Drymanthele actylis glomerata Sesleria argentea Puccinellia distans Poa infirma Secale cereale 1 distachyon 0.1 Figur Combined ITS/trnL-F: Bayesian 50% MR consensus tree of 9643 trees. Posterior probability 5 is nic М 100 on the corresponding branches. Species numbers correspond to different accessions of the same taxon (se Table 145 Phylogenetic Relationships of Мира Torrecilla et al. 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