A Revision of Trisetum (Poaceae: Pooideae Aveninae) in South America

Victor Finot

Ribosomal DNA sequences were employed to infer relationships among EuropeanPottiaceae. Intragenic spacer I was sequenced for sixTrichostomoideae (Eucladium verticillatum, Pleurochaete squarrosa, Tortella flavovirens, T. nitida, T. tortuosa, Trichostomum brachydontium) and seven taxa from the other European subfamilies included inPottiaceae (Bryoerythrophyllum recurvirostrum, Didymodon vinealis, Microbryum starkeanum, Tortula muralis, Syntrichia ruralis, Weissia controversa andTimmiella cf.barbuloides). Cladistic analysis of sequence data shows close relationships betweenPleurochaete, Tortella andWeissia. Tortella appears to be paraphyletic, as it includesPleurochaete, Weissia andTrichostomum. Weissia, therefore, seems to be better placed inTrichostomoideae than, as traditionally done, inPottioideae. NeitherMerceyoideae norPottioideae appear to be monophyletic sensu stricto, at least within the limits of the taxa in analysis. Within the latter subfamily,Syntrichia is clearly separated fromTortula. These results would suggest that taxonomy inPottiaceae does not depict the pattern of descent, and therefore is in need of revision.

Victor L. Finot,' Paul M. Peterson,3 Fernando 0 Zuloaga,* Robert J. sorene, and Oscar Mattnei A REVISION OF TRISETUM (POACEAE: POOIDEAE: v AVENINAE) IN SOUTH AMERICA1 ABSTRACT A taxonomic treatment of Trisetum Pers. for South America, is given. Eighteen species and six varieties of Trisetum are recognized in South America. Chile (14 species, 3 varieties) and Argentina (12 species, 5 varieties) have the greatest number of taxa in the genus. Two varieties, T. barbinode var. sclerophyllum and T longiglume var. glabratum, are endemic to Argentina, whereas T. mattheii and T nancaguense are known only from Chile. Trisetum andinum is endemic to Ecuador, T. macbridei is endemic to Peru, and T. foliosum is endemic to Venezuela. A total of four species are found in Ecuador and Peru, and there are two species in Venezuela and Colombia. The following new species are described and illustrated: Trisetum mattheii Finot and T nancaguense Finot, from Chile, and T pyramidatum LouisMarie ex Finot, from Chile and Argentina. The following two new combinations are made: T barbinode var. sclerophyllum (Hack, ex Stuck.) Finot and T. spicatum var. cumingii (Nees ex Steud.) Finot. A key for distinguishing the species and varieties of Trisetum in South America is given. The names Koeleria cumingii Nees ex Steud., Trisetum sect. Anaulacoa Louis-Marie, Trisetum sect. Aulacoa Louis-Marie, Trisetum subg. Heterolytrum Louis-Marie, Trisetum subg. Isolytrum Louis-Marie, Trisetum subsect. Koeleriformia Louis-Marie, Trisetum subsect. Sphenopholidea Louis-Marie, Trisetum malacophyllum Steud., Trisetum variabile E. Desv., and Trisetum variabile var. virescens E. Desv. are lectotypified. Key words: Aveninae, Gramineae, Poaceae, Pooideae, Trisetum. RESUMEN Se realizd un estudio taxonomico del genero Trisetum en America del Sur. Se reconocio 18 especies y 6 variedades sudamericanas de Trisetum. Chile (14 especies, 3 variedades) y Argentina (12 especies, 5 variedades) poseen el mayor niimero de taxones del genero Trisetum. Dos variedades, T. barbinode var. sclerophyllum y T. longiglume var. glabratum. son endemicos de Argentina, mientras T. mattheii y T. nancaguense se conocen solo para Chile. Cuatro especies se encuentran en Ecuador y Peru y dos especies en Venezuela y Colombia. Trisetum andinum es endemica del Ecuador, T macbridei es endemica del Peru, y T foliosum es endemica de Venezuela. Se describen e ilustran las siguientes nuevas especies: Trisetum mattheii Finot y T nancaguense Finot, de Chile, y T. pyramidatum Louis-Marie ex Finot, de Chile y Argentina. Se establecen las siguientes dos nuevas combinaciones: T barbinode var. sclerophyllum (Hack, ex Stuck.) Finot y T spicatum var. cumingii (Nees ex Steud.) Finot. Se entrega una clave para separar las especies y variedades de Trisetum en Sudamerica. Los nombres Koeleria cumingii Nees ex Steud., Trisetum sect. Anaulacoa LouisMarie, Trisetum sect. Aulacoa Louis-Marie, Trisetum subg. Heterolytrum Louis-Marie, Trisetum subg. Isolytrum. LouisMarie, Trisetum subsect. Koeleriformia Louis-Marie, Trisetum subsect. Sphenopholidea Louis-Marie, Trisetum malacophyllum Steud., Trisetum variabile E. Desv., y Trisetum variabile var. virescens E. Desv. fueron lectotipificados. The genus Trisetum was described by Persoon (1805), including 11 species previously treated un- habiting temperate and cold zones in both hemispheres (Louis Marie, 1928, 1929; Swallen, der Avena L. As currently defined, Trisetum com- 1948; Tsvelev, 1970, 1983; Jonsell, 1980; Veld- prises about 70 species of perennial grasses, in- kamp & Van der Have, 1983; Clayton & Renvoize, 1 This paper is part of the doctoral thesis of VLF in the Dpto. Botanica, Universidad de Concepcion, Concepcion, Chile. We thank Vladimiro Dudas for preparing the illustrations. Thanks are due to the Directors and Curators of the following herbaria: BA, BAA, BAF, CONC, CTES, CR, F, LP, MERL, P, QCA, S, SI, US, and ZOELLNER. We thank Silvia Denham, Victoria C. Hollowell, Osvaldo Morrone, Dan Nicolson, and an anonymous reviewer for suggesting improvements to the manuscript. VLF greatfully acknowledges the Myndel Foundation for a fellowship to study types of Trisetum at Paris (P) and Stockholm (S); the MINEDUC for a fellowship to study types at the Smithsonian Institution, Washington and the Instituto de Rotanica Darwinion, Buenos Aires; the financial support from Direccidn de Investigacion, Universidad de Concepcion, Project DIUC 201.121.005-1.0 and 204.121.009-1.0; and OM and Clodomiro Marticorena (U. de Concepcion, Chile) for helping direct my Ph.D. 2 Universidad de Concepcion, Facultad de Agronomia, Casilla 537, Chilian, Chile. vifinot@udec.cl. 3 Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, U.S.A. peterson@si.edu; sorengr@si.edu. 4 Instituto de Botanica Darwinion, Labarden 200, San Isidro, Argentina, fzuloaga@darwin.edu.ar. ANN. MISSOURI BOT. GARD. 92: 533-568. PURLISHED ON 28 DECEMBER 2005. 534 Annals of the Missouri Botanical Garden 1986; Pohl & Davidse, 1994; Tucker 1996; Edgar, 1998; Finot et al., 2004). In America, the genus is distributed from Greenland to southern South America, ranging from approximately 69°N to 55°S (Hulten, 1959; Nicora, 1978). Trisetum has been traditionally divided in two sections: section Trisetum, typified by T. flavescens (L.) P. Beauv., with lax, open panicles and culms glabrous below the inflorescence, and section Trisetaera Asch. & Graebn., typified by T. spicatum (L.) K. Richt., with dense, spiciform panicles and culms pilose below the inflorescence. This supraspecific classification has been accepted by most authors (Chrtek, 1965; Chrtek & Jirasek, 1963; Tsvelev, 1983). Chrtek (1965) divided Trisetum in four subgenera: subgenus Trisetum, subgenus Distichotrisetum Chrtek, subgenus Glaciotrisetum Chrtek, and subgenus Graciliotrisetum Chrtek and divided subgenus Trisetum into five sections: section Trisetum, section Trisetaera Asch. & Graebn., section Rigida Chrtek, section Hispanica Chrtek, and section Carpatica Chrtek. Only the Trisetum sections Trisetum and Trisetaera include American representatives. A general account of the American taxa was first made by Steudel (1853•1855), who described ten species for South America: Trisetum andinum Benth., T barbatum Steud., T hirtum Trin., T caudulatum Trin., T barbinode Trin., T splendidulum Steud., T airiforme Steud., T heteronymum Steud., T malacophyllum Steud., and T phleoides (d'Urv.) Kunth. Hitchcock (1927) later recognized the following six species from Ecuador, Peru, and Bolivia: Trisetum deyeuxioides (Kunth) Kunth, T scabrifolium Hitchc, T floribundum Pilg., T macbridei Hitchc, T spicatum (L.) K. Richt., and T andinum Benth. Louis-Marie (1928, 1929), in his taxonomic revision of Trisetum, recognized 60 species for America. However, Louis-Marie had a very broad concept of Trisetum that included several species later transferred to other genera, including Bromus, Deschampsia, Dielsiochloa, Giaphephorum, Leptophyllochloa, Peyritschia, and Sphenopholis (Valencia, 1941; Pilger, 1943; Parodi, 1949a; Erdman, 1965; Nicora, 1978; Finot, 2003; Finot et al., 2004). Louis-Marie (1928, 1929) divided the genus Trisetum into two subgenera (Table 1; see Appendix 3 for subgeneric lectotypifications): subgenus Heterolytrum Louis-Marie and subgenus Isolytrum LouisMarie. He divided Trisetum subg. Heterolytrum into two sections: section Anaulacoa Louis-Marie (including subsect. Trisetum, subsect. Sphenopholidea Louis-Marie, subsect. Giaphephorum (Desv.) LouisMarie, subsect. Koeleiiformia Louis-Marie, and subsect. Deschampsioidea Louis-Marie) and section Aulacoa Louis-Marie. South American species in Trisetum sect. Anaulacoa subsect. Trisetum included: T. splendidulum, T irazuense (Kuntze) Hitchc, T fraudulentum Steud., T. malacophyllum Steud., T cernuum Trin., T. heteronymum Steud., T. hirtum Trin., T mollifolium Louis-Marie, T oreophilum Louis-Marie var. oreophilum, T oreophilum var. johnstonii Louis-Marie, T lasiolepis E. Desv., T preslei (Kunth) E. Desv., T spicatum var. spicatum, T spicatum var. hirsutum Louis-Marie, T. spicatum var. phleoides (Kunth) Macloskie, T spicatum var. fuegianum (Hack.) Louis-Marie, T spicatum var. dianthemum Louis-Marie, T. spicatum var. andinum (Benth.) Louis-Marie, T barbinode Trin. var. barbinode, T. barbinode var. hirtiflorum (Hack.) LouisMarie, T caudulatum Trin., T variabile E. Desv. var. variabile, T variabile var. flavescens E. Desv., T variabile var. virescens E. Desv., T variabile var. chiloense (Phil.) Louis-Marie, T variabile var. vidalii (Phil.) Louis-Marie, T erectum Phil., T monticola Phil., and T paradoxum Phil. In Trisetum subsect. Koeleiiformia Louis-Marie included the following seven species: T. laxiflorum Phil., T araeanthum Phil., T brachyatherum Phil., T. depauperatum Phil., T. micratherum E. Desv., T. nemorosum Phil., and T laxum Phil. Trisetum micratherum was later transferred to Leptophyilochloa by Calderon (1978) (L. micrathera (E. Desv.) C. E. Calderon). Tiisetum subsect. Sphenopholidea and subsection Giaphephorum do not have South American representatives. Louis-Marie included two South American species: T brasiliense Louis-Marie and T juergensii Hack., in Tiisetum subsect. Deschampsioideum. In Tiisetum sect. Aulacoa LouisMarie included two species, Bromus trinii E. Desv. (replaced name Tiisetum hirtum Trin. = Bromus berteroanus Colla) and T. floribundum Pilg. (= Dielsiochloa floribunda (Pilg.) Pilg.). South American representatives of Tiisetum subg. Isolytrum included T deyeuxioides, T. longiglume Hack., T andicola Louis-Marie, T. evolutum (E. Fourn.) Hitchc, and T macbridei Hitchc. Valencia (1941) placed Trisetum juergensii, T confertum Pilg., T brasiliense, and T. andicola in Deschampsia. However, Parodi (1949b) disagreed with Valencia concerning the position of T andicola. According to Parodi (1949b), a lemma with two apical setae produced by the apical extension of the intermediate nerves supports the recognition of this species in genus Trisetum. Hulten (1959) reviewed the taxonomy of the Trisetum spicatum complex on a world-wide basis and he cited several South American taxa, including a new subspecies, T spicatum subsp. bolivianum Volume 92, Number 4 2005 Finot et al. Trisetum in South America 535 Hulten from La Paz, Bolivia. Hulten did not treat most of the varieties of T. spicatum s.l. mentioned by Louis-Marie for South America. In addition to the European T. flavescens introduced in North and South America, Nicora (1978) recognized 13 species in Patagonia: T. cernuum, T. longiglume Hack., T. tomentosum (E. Desv.) Nicora, T. spicatum, T. phleoides (d'Urv.) Kunth, T. cumingii (Nees ex Steud.) Nicora, T. lechleri (Steud.) Nicora, T. caudulatum, T. barbinode, T. hirtiflorum Hack., T. sclerophyllum Hack., T. lasiolepis E. Desv., and T. preslei (Kunth) E. Desv. Later, Rugolo de Agrasar and Nicora (1988) described a new species, T. ambiguum Rugolo & Nicora, from Argentina. Several species are controversial in the generic alignment. Although Trisetum subsect. Deschampsioideum is retained for several species of Mexico and Central America (Finot et al., 2004), T. brasiliense and T. juergensii of South America do not fit well in the genus and will be dealt with in future papers. In addition, we recognize Trisetum confertum as Peyritschia conferta (Pilg.) Finot (Finot, 2003). Our current treatment of Trisetum for the Americas recognizes 45 species and 20 varieties (Finot, 2003; Finot et al., 2004; this paper). In this paper we recognize 18 species and six varieties for South America (see Appendix 1 for alphabetical list). We describe and illustrate three new species of Trisetum (T. mattheii Finot, T. nancaguense Finot, and T. pyramidatum Louis-Marie ex Finot), make two new combinations (T. barbinode var. sclerophyllum (Hack, ex Stuck.) Finot and T. spicatum var. cumingii (Nees ex Steud.) Finot), and provide descriptions and keys to the species and varieties in South America. MATERIAL AND METHODS Type specimens and general collections from the following herbaria were studied: BA, BAA, BAF, CONC, CTES, CR, F, LP, MERL, P, QCA, S, SI, US, and ZOELLNER. In some cases, the curators of these herbaria sent us digital photographs or Xerox copies of types, or we had access to internet digital images (C, PR, NY). Abbreviations for herbaria correspond to those cited by Holmgren et al. (1990), except ZOELLNER, which is the personal herbarium of Otto Zoellner, Quilpue, Chile. An index to specimens examined and a list of all names and synonyms mentioned in this manuscript, including those in the introduction, are treated in Appendix 2 and 4, respectively. In the morphological descriptions the length given for florets was usually taken from the first or lowest floret. If there were three or more florets per spikelet, then the second floret was sometimes used to calculate the range. Therefore, when using our keys to determine South American specimens of Trisetum it is best to measure only the first or lowest floret. KEY TO TRISETUM AND MORPHOLOGICALLY SIMILAR GENERA IN SOUTH AMERICA la. Plants annual. 2a. Plants 20•250 cm tall; panicles lax, open; spikelets 15•50 mm long, pendulous; glumes equal in length; lemmas dorsally awned, the awn geniculate and bent at the base; ovary pilose; caryopsis with solid endosperm Avena 2b. Plants 3•50 cm tall; panicles dense, spiciform; spikelets 2.8•4.5(•7) mm long; glumes unequal in length; lemmas muticous or with a short subapical awn up to 3 mm long; ovary glabrous; caryopsis with liquid endosperm Rostraria lb. Plants perennial, caespitose, rhizomatous or with bulbous bases. 3a. Spikelets 7- to 9-flowered, with 2 or 3 fertile basal florets and 4 to 6 sterile upper florets; rachilla disarticulating only immediately above the glumes, florets falling as a collective unit Dielsiochloa 3b. Spikelets (1)2- to many-flowered, without sterile florets or the terminal floret reduced; rachilla disarticulating above the glumes and between the florets (except Arrhenatherum where 2 florets fall together). 4a. Lemmas muticous or with a small awn inserted near the apex, the aw7n straight. 5a. Panicles lax, open; lemmas 3-nerved Leptophyllochloa 5b. Panicles contracted, subspiciform; lemmas 5-nerved. 6a. Lemmas strongly scabrous, the keel w7ith shining ciliate hairs; florets w7ith a single stamen Raimundochloa 6b. Lemmas glabrous, the keel scabrous, rarely ciliate (ciliate in Koeleria fueguina C. E. 7 Calderon ex Nicora); florets w ith 3 stamens. 7a. Spikelets 2- or 3(4)-flowered; ovary glabrous or with a few short hairs near the apex; caryopsis fusiform, without a ventral groove, hilum punctiform, endosperm liquid Koeleria 7b. Spikelets (l)2(3)-flowered; ovary pilose; caryopsis sub-triangular, with a ventral groove, hilum linear, 1/5•1/3 as long as the caryopsis; endosperm solid ___ Relchela 4b. Lemma with a dorsal awn inserted well below the apex, the awn geniculate and bent. 8a. Spikelets 2-flowered; the low7er floret staminate with a large geniculate awn inserted on the 536 Annals of the Missouri Botanical Garden Table 1. Infrageneric classification of the South American species of Trisetum in Louis-Marie's (1928, 1929) and our taxonomic treatment. Accepted taxa are presented in bold. Louis-Marie (1928, 1929) Genus Trisetum Subg. Heterolytrum Sect. Anaulacoa Subsect. Trisetum (as Eutriseta) T. cernuum Trin., T. fraudulentum Steud. T. irazuense (Kuntze) Hitchc. T. flavescens (L.) P. Beauv. T spicatum var. andinum (Benth.) LouisMarie T. barbinode Trin. var. barbinode T. barbinode var. hirtiflorum (Hack.) Louis-Marie T. caudulatum Trin., T splendidulum Steud., T heteronymum Steud., T. variabile Yax. flavescens E. Desv., T variabile var. virescens E. Desv., T variabile var. chiloense (Phil.) Louis-Marie, T variabile var. vidalii (Phil.) Louis-Marie, T. monticola Phil. T spicatum var. dianthemum Louis-Ma T. oreophilum var. jolinstonii LouisMarie T. oreophilum Louis-Marie var. oreophilum T spicatum var. hirsutum Louis-Marie, T spicatum var. phleoides (d'Urv.) Macloskie T. preslei (Kunth) E. Desv., T lasiolepis E. Desv. T. spicatum (L.) K. Richt., T. spicatum var. fuegianum (Hack.) Louis-Marie T malacophyllum Steud., T. mollifolium Louis-Marie T hirtum Trin. Trisetum erectum Phil. T paradoxum Phil. Subsect. Koeleriformia T laxiflorum Phil., T araeanthum Phil., T brachyatherum Phil., T depauperatum Phil., T nemorosum Phil., T. laxum Phil. This treatment Genus Trisetum Subg. Trisetum (Subg. Trisetum) (Subg. Trisetum sect. Trisetum) Sect. Trisetum T. cernuum T. irazuense T. flavescens T. foliosum Swallen Sect. Trisaetera Asch. & Graebn. T. ambiguum Rugolo & Nicora T. andinum Benth. T. barbinode var. barbinode T. barbinode var. hirtiflorum T. barbinode var. sclerophyllum (Hack.) Finot T. caudulatum Trin. var. caudulatum T. caudulatum Trin. var. correae Nicora T. dianthemum (Louis-Marie) Finot T. longiglume Hack. T. longiglume var. glabratum Nicora T. macbridei Hitchc. T. mattheii Finot T. nancaguense Finot T. oreophilum var. jolinstonii T. oreophilum var. oreophilum T. phleoides (d'Urv.) Kunth T. preslei T. pyramidatum Louis-Marie ex Finot T. spicatum T. spicatum var. cumingii (Nees ex Steud.) Finot Bromus sect. Neobromus (Shear) Hitchc. B. berteroanus Colla Unknown status Unknown status Leptophyllochloa C. E. Calderdn L. micranthera (E. Desv.) C. E. Calderon Volume 92, Number 4 2005 Table 1. Finot et al. Trisetum in South America 537 Continued. Louis-Marie (1928, 1929) Subsect. Deschampsioidea Louis-Marie Trisetum brasiliense Louis-Marie Subsect. Sphenopholidea Louis-Marie (without representatives in South America) Subsect. Graphephorum (Desv.) Louis-Marie (without representatives in South America) Sect. Aulacoa T trinii (E. Desv.) Louis-Marie T floribundum Pilg. Subg. Isolytrum T deyeuxioides (Kunth) Kunth, T. evolutum (E. Fourn.) Hitchc. T. longiglume Hack., T andicola LouisMarie T. macbrldei Hitchc. This treatment Subg. Deschampsioideum (Louis-Marie) Finot (without representatives in South America) Deschampsia brasiliensis (Louis-Marie) Valencia Sphenopholis Scribn. (without representatives in South America) Graphephorum Desv. (without representatives in South America) Dielsiochloa Pilg. Bromus berteroanus Colla Dielsiochloa floribunda (Pilg.) Pilg. Peyritscliia deyeuxioides (Kunth) Finot see T. sect. Trisaetera see T. sect. Trisaetera lower 1/3 of the lemma; upper floret perfect, the apex muticous or with a small subapical awn Arrhenatherum 8b. Spikelets (1)2- to many-flowered; florets all perfect. 9a. Spikelets 3- to 8-flowered; lemmas 7- to 9-nerved; panicles terminal and with cleistogamous axillary panicles Amphibromus 9b. Spikelets (1)2- or 3(to 6)-flowered; lemmas 5-nerved; panicles terminal and, if present, axillary panicles not cleistogamous. 10a. Culms with a bulbous base; panicles spiciform; glumes shorter than all florets (together); lemmas rounded on the back, the awn inserted near the middle; lemma apex bilobed, hyaline; callus acute Helictotrichon 10b. Culms without a bulbous base; panicles open or spiciform; glumes shorter to longer than all florets; lemma keeled, somewhat laterally flattened, the awn inserted on the upper 1/3; lemma apex bidentate with intermediate nerves prolonged into setae; callus obtuse Trisetum TAXONOMIC TREATMENT Trisetum Pers., Syn. PL 1: 97. 1805. TYPE: Trisetum flavescens (L.) P. Beauv., Ess. Agrostogr. 88, 153, t. 18, f. 1. 1812, lectotype, designated by Hitchcock, U.S.D.A. Bull. 772: 107-109. 1920. Rupestrina Prov., Fl. Canad.: 689. 1862. TYPE: Rupestrina pubescens Prov. (= Trisetum spicatum (L.) Richt.) Perennials, caespitose, sometimes shortly rhizomatous; culms 5•300 cm tall, erect to geniculate at base, glabrous or pubescent. Leaf sheaths glabrous or pubescent, longer or shorter than the internocles; blades flat, conduplicate, convolute or involute, soft, rarely rigid; ligule membranous. Inflorescence in panicles contracted or open, spiciform, ovate or pyramidal; the rachis glabrous, scabrous, or pubescent. Spikelets (1)2- to 6-flowered, short pedicellate; rachilla pubescent or glabrous, 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 1- to 3-veined, usually shorter and narrower than the second, second glume 3- to 5-nerved; lemmas lanceolate, (3 to)5(to 7)-veined, usually awned or muticous, 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 1/3, rarely the middle, of the subapical portion of the lemma; awn exerted, geniculate or merely divaricate; callus short pilose; palea not tightly enclosed by the margins of the lemma (gaping), 2-keeled, hyaline, usually shorter than the lemma; stamens 3, anthers 0.3•4.5 mm long; lodicules 2, membranous; ovary glabrous or with short and shining trichomes near the apex. Caryopses compressed, soft; hilum short, punctiform; endosperm solid or liquid, soft or hard. Basic chromosome number x = 7. Comments. Our subgeneric treatment of the 538 Annals of the Missouri Botanical Garden species that occur in South America includes a sin- numbered 1•4 are placed in Trisetum sect. Trisetum gle subgenus: Trisetum subg. Trisetum with two sec- and species numbered 5•18 are placed in Trisetum tions: section Trisetum and section Trisetaera. In the sect. Trisetaera. A description of Trisetum subg. Tri- following key we have indicated in which section setum and Trisetum sects. Trisetum and Trisetaera each species resides (see leads la and lb). Species appears in Finot et al. (2004). KEY TO SPECIES AND VARIETIES OF TRISETUM IN SOUTH AMERICA la. Panicles lax, mostly open or narrow, pyramidal, never spiciform; culms glabrous below the inflorescence; glumes unequal, the first glume shorter and narrower than the second glume; glumes notably shorter than the florets (Trisetum sect. Trisetum). 2a. Sheath apex notably extended upward on one side as an appendix as long as the ligule 3. T. foliosum 2b. Sheath apex not extended as an appendix. 3a. Ovary and caryopsis hairy at the apex; panicles few-flowered, the lower branches usually naked below; first glume 0.5•5 mm long, sometimes reduced; ligule 1.5•7 mm long 1. T. cernuum 3b. Ovary and caryopsis glabrous at the apex; panicles densely-flowered, branches not naked below; first glume 2•5 mm long, never reduced. 4a. Panicles usually gold-yellowish; lemmas not strongly scabrous; ligule 0.5•2 mm long; leaf blades (3•) 10•16 cm long, 2^4 mm wide 2. T. flavescens 4b. Panicles greenish to purplish; lemmas strongly scabrous; ligule 1•^ mm long; leaf blades 20•30 cm long, 2.5•6 mm wide 4. T. irazuense lb. Panicles spiciform to contracted, never open; culms pubescent or hairy below the inflorescence, rarely glabrous to subglabrous; glumes subequal, a little shorter, equal or longer than the florets (Trisetum sect. Trisetaera). 5a. Lemma hairy, the hairs usually more than 0.5 mm long. 6a. Plants small, usually less than 20 cm tall; panicle dense, spiciform. 7a. Leaf glabrous; glumes equaling or slightly exceeding the florets; first glume 4.5•6 mm long; second glume (4.5•)5.5•6.6 mm long 16. T. preslei 7b. Leaf hairy; glumes shorter than the florets; first glume 2.7•4 mm long; second glume 3.3• 4.7 mm long 14b. T. oreophilum var. johnsionii 6b. Plants taller, usually more than 30 cm tall; panicle spiciform or contracted. 8a. Glumes longer or equaling the florets, the second glume usually longer than the florets; first glume 1- or 3-nerved; leaf glabrous. 9a. Panicles contracted, pale green, up to 15 cm long; glumes unequal; second glume 6.5•10 mm long; leaf blades soft 7a. T. barbinode var. barbinode 9b. Panicles spiciform, gold-purple, (2•)3•5(•8) cm long; glumes subequal to unequal; second glume 5•8 mm long. 10a. Ovary glabrous; blades stiff 7c. T. barbinode var. sclerophyllum 10b. Ovary with short trichomes on the apex; blades soft 7b. T. barbinode var. hirtiflorum 8b. Glumes shorter than the florets; first glume 1-nerved; leaf blades pubescent. 11a. Panicles silvery-green to weakly purple, not purplish; spikelets 5.5•8 mm long; first glume 3.5•6 mm long; second glume 5.3•7.6 mm long 13. T. nancaguense lib. Panicles green-purplish to strongly purplish; spikelets 3.5•5 mm long; first glume 2.2• 4.5 mm long; second glume 2.8•4.7 mm long. 12a. Panicles 0.8•1.5 cm wide, dense, many-flowered; spikelets 3.5•4.5 mm long; first glume 2.2•3.6 mm long; second glume 2.8•4.1 mm long 14a. T. oreophilum var. oreophilum 12b. Panicles 0.5•0.8 cm wide, narrow, few-flowered; spikelets 4.5•5 mm long; first glume 3.5•4.5 mm long; second glume 3.8^4.7 mm long 12. T. mattheii 5b. Lemma glabrous or scabrous, never hairy, but rarely covered with very short hairs, the hairs shorter than 0.5 mm long. 13a. Panicles no more than 3 to 4 times as long as wide, dark purplish; plants velvety; lemma scabrous or covered with very short trichomes giving a velvety texture 6. T. andinum 13b. Panicles more than 3 to 4 times as long as wide, green to more or less purplish; plants glabrous or pilose, not velvety; lemmas glabrous or scabrous, not velvety 14a. Glumes longer than the florets, subequal in length and width. 15a. Plant totally pubescent; glumes ciliate on the keel; first glume 4•6.5 mm long, second glume 5•6.5 mm long 15. T. phleoides 15b. Plants not totally pubescent; glumes scabrous on the keel, not ciliate; glumes more than 6•9.5 mm long. 16a. Glumes 7.5•8 mm long, ovate, exceeding the florets by 1/3•1/2 in length; blades pilose 11. 73 macbridei 16b. Glumes 6•9.5 mm long, lanceolate, exceeding the florets by 1/4•1/3 in length; blades glabrous. Volume 92, Number 4 2005 Finot et al. Trisetum in South America 539 17a. Spikelets 6.5•8 mm long; awn borne on the upper 1/3 of the lemma; callus trichomes short (ca. 0.2 mm long); ovary glabrous at the apex 9. T. dianthemum 17b. Spikelets 9•10 mm long; awn borne on the middle of the lemma; callus trichomes long, reaching 1/2•3/4 the length of the lemma; ovary with or without trichomes at the apex. 18a. Panicles ovoid, dense, subspiciform, many-flowered; ovary and caryopsis with short, curved and shining trichomes near the apex; caryopsis 2•2.7 mm long; anthers (0.5•)0.8•1 mm long 10a. T. longiglume var. longiglume 18b. Panicles linear, narrow, few-flowered; ovary and caryopsis glabrous near the apex; caryopsis 3.5•4.2 mm long; anthers 1.3•1.7 mm long 10b. T. longiglume var. glabratum 14b. Glumes equal or shorter than the florets, dissimilar in length and width, or the first glume shorter and the second glume longer than the florets and first glume shorter and narrower than the second glume. 19a. Spikelets 1-flowered; ovary with a few trichomes at the apex; lemma conspicuously nerved toward the apex, the intermediate and marginal nerves prolonged beyond the apex as four short apical awns 5. T. ambiguum 19b. Spikelets 2- or 3(to 5)-flowered; ovary glabrous at the apex; lemma inconspicuously nerved toward the apex, the intermediate nerves prolonged beyond the apex as 2 setae or short apical awns, the marginal nerves not prolonged beyond the apex. 20a. Second glume longer than the spikelet, the first glume shorter than the spikelet. 21a. Plants rhizomatous; panicle pyramidal, contracted, not spiciform, 7•11 cm long, 2•3 cm wide; paleas always shorter than the lemma 17. T. pyramidatum 21b. Plants caespitose; panicle spiciform, dense, 4•10 cm long, 0.8•2 cm wide; paleas a little shorter than the lemma in the first floret and longer than the lemma in the upper floret 18b. T. spicatum var. cumingii 20b. Glumes (second and first) shorter than the florets. 22a. Glumes similar in width, the first glume 0.5•1 mm wide, first glume as wide as second glume or a little narrower; culms densely pubescent below the panicles; blades glabrous or pilose; spikelets 2- or 3-flowered 18a. T. spicatum var. spicatum 22b. Glumes dissimilar in width, the first glume 0.3•0.5 mm wide, narrower than the second glume; culm glabrous to subglabrous below the panicles: blades pilose; spikelets 2- to 4(5)-flowered. 23a. Ovary and caryopsis glabrous at the apex 8a. T. caudulatum var. caudulatum 23b. Ovary and caryopsis with trichomes at the apex 8b. T. caudulatum var. correae Trisetum subg. Trisetum sect. Trisetum sheaths longer or shorter than the internodes, gla- 1. Trisetum cernuum Trin., Mem. Acad. Imp. brous or more rarely sparsely pilose; ligule 1.5•7 Sci. St. Petersbourg Ser. 6, Sci. Math. 1 (1): mm long, longer in the upper leaves, oval, fimbri- 61. 1830. Avena cernua (Trin.) Kunth, Rev. ate, ciliate on the margins, the dorsal surface gla- Gen. 1, suppl. 26. 306. 1833. TYPE: U.S.A. brous or more frequently pilose; blades 10•22 X Alaska, Sitka, J. F. G. von Eschscholtz s.n. (ho- 0.3-1.2 cm, flat, soft, glabrous or sparsely pubes- lotype, LE not seen; isotypes, BAA-3366!, P!, cent adaxially. Panicles 6•30 X 2•5 cm, lax, open, US-81779!). Avena leptostachys Hook, f., Fl. Antarct.: 378. 1846. TYPE: Chile. Strait of Magalhaens, Port Famine, ,••, • •. ,, , • . 18ZO, Lapt. King s.n. (nolotype, K not seen; isotypes, SCO ex K!, US-fragm. ex K!). Trisetum fraudulentum Steud., Syn. PL Glumac. 1: 424. 1854. TYPE: Chile. Sandy Point, Magellan, W. Lechlev 1283 (holotype, P!; isotypes CONC fragm. ex P!, ITC Q^Q/IQO fragm. A.••-~ ex •• LE!, TT7I US ITC fragm. f ~• P-STEUDU CTT7ITr» m US-868480 ex 434 & photo!). nodding, green or purple; the branches capillary, flexuous, the lower branches usually naked toward the base or with spikelets to the base. Spikelets . r -.^ ., r, A /<-»\I A n 1 i- 1 n 4.5•Iz X Z-^i mm, (zloor 4-nowered; r pedicels 1• v ' 4.5 mm long, capillary, flexuous, glabrous or scabrous; rachilla 1.5-2 mm long, pilose with stiff trichomes 1-2 mm long; glumes shorter than the y2 t 2/g , fl t th ikelet un_ ' « V ' J equal to subequal in length and very unequal in width, the first glume narrower than the second; Perennial, with short rhizomes; culms 35•85 cm keel smooth to scaberulous; margins hyaline, some- tall, glabrous, up to 3 mm diam. near base, some- times scabrous-ciliate; first glume 0.5•5 X 0.1•0.5 what geniculate; nodes 2 to 4, glabrous. Leaf mm, usually 1/2•2/3 as long as the second glume, 540 Annals of the Missouri Botanical Garden linear-lanceolate, attenuate, 1-nerved, sometimes reduced; second glume 3•7.5 X 0.5•1 mm, ovallanceolate, 3-nerved; florets 5•8.5 X 0.7•1 mm; lemmas linear-lanceolate, glabrous, scabrous toward the apex, awned, green or tinged with purple on the margins and apex; apex 2-aristulate, the aristae 0.5•1.5 mm long; awn 6•16 mm long, borne on the upper 1/3 ca. 1.5•2.5 mm below the apex, curved, not strongly twisted nor geniculate, scabrous, up to 3 times as long as the lemma; callus obtuse, pubescent, the trichomes 0.2•0.7 mm long; paleas 3.5•5.7 mm long, shorter than the lemma, hyaline, 2-nerved, the nerves scabrous; apex bidentate; anthers 0.5•1.5 mm long; lodicules 0.7• 0.8 mm long; apex bilobed; ovary with short and curved trichomes near the apex. Caryopses 2.5 X 1 mm, hairy at the apex; endosperm soft. first glume. The first glume, usually 2•3.8 mm long, can be reduced to 0.5 mm long. However, we found spikelets with reduced and non reduced glumes in the same plants (e.g., Bolander 6122, collected in California, U.S.A.). Trisetum cernuum is morphologically similar to T flavescens and T. irazuense. Trisetum cernuum differs from T. flavescens in having paucispiculate panicles, hairy caryopses, and shorter glumes that are sometimes reduced. Trisetum flavescens has multispiculate panicles, glumes that are never reduced, and caryopses that are glabrous at the apex. Trisetum flavescens is a European species, introduced in southern South America as a forage plant (Nicora, 1978). Trisetum irazuense differs from T. cernuum in having lemmas strongly scabrous and the ovary glabrous at the apex. It shares with T cernuum lax, open panicles, spikelets with glumes shorter than the florets, and dissimilar glumes that are never reduced. Trisetum irazuense is distributed in Mexico, Costa Rica, Panama, Honduras, Colombia, Venezuela, Ecuador, and Peru (Hitchcock, 1927; Pohl, 1980; Hernandez-Torres & Koch, 1988; Davidse et al., 1994; J0rgensen & Ulloa, 1994; Finot et al., 2004). Distribution and habitat. Disjunct,, occurring in western North America, west of the Rocky Mountains, from southwestern Alaska to northern California, and in southern South America. In South America, Trisetum cernuum occurs in Patagonia of Argentina and Chile usually below 2000 m, from 38°S to approximately 55°S, south of Estrecho de Magallanes. In Chile, it grows from the Region IX (Malleco, ca. 38°S) to the Region XII (Isla Navarino, ca. 54°55'S). In Argentina, T. cernuum is found in Chubut, Neuquen, Rio Negro, and Tierra del Fuego (Nicora, 1978; Zuloaga et al., 1994), from 38°S in Chubut to 54°47'S in Ushuaia. This species is frequently associated with moist woods of Araucaria araucana (Mol.) K. K.och-Nothqfagu.s dombeyi (Mirb.) Oerst. and Nothqfagus pumilio (Poepp. & Endl.) Krasser, and "marlines" forests between sea level and 1500 m. Phenology. Flowering between January and March. Illustrations. Hitchcock and Chase (1950: 289, fig. 389); Nicora (1978: 242, fig. 156 A-D). Comments. Trisetum cernuum is a well-defined species, easily recognized by its lax, open, nutant panicles with subverticillate and paucispiculate branches, the lower ones without spikelets below the middle, spikelets with dissimilar glumes, glumes shorter than the spikelets, and caryopses with short hairs at the apex. These hairs, very typical in the genus Trisetum, are also present in other South American species such as T ambiguum, T caudulatum var. correae, T. barbinode var. hirtiflorum, and T. longiglume var. longiglume. All these species are restricted to Chile and Argentina, and they are easily distinguished from T cernuum by their spiciform panicles. Trisetum cernuum shows extreme variation in glume length, especially in the Additional specimens studied. ARGENTINA. Chubut: Lago Fontana, 10 Feb. 1932, Castellanos s.n. (BA, S). Neuquen: Los Lagos, Villa La Angostura, Pedersen 1523 (CTES). Rio Negro: Bariloche, Valle def Chollhuaco, Refugio Neumeyer, Riigolo de Agrasar et al. 12381 (CONC); Region montafiosa cercana al Lago Nahuel Huapi', Parodi 11827 (BAA); P.N. Nahuel Huapi, Cerro Lopez, Perez Moreau 1949 (BA); Lago Nahuel Huapi, Cerro Catedral, Cabrera 11511 (LP). Tierra del Fuego: Bahia Lapataia, 2 Feb. 1948, Perez Moreau & Guarrera s.n. (BA); Valle de Tierra Mayor, Ruiz Leal & Roig 15023 (MERL, BAA); Ushuaia, source gauche du Rio Grande, 7 Mar. 1896, Alboff s.n. (CORD); Ushuaia, cerros alrededores de Ushuaia, Grondona 5695 (BAA). CHILE. IX Region: Malleco, Cordillera de Las Raices, Matthei & Bustos 111 (CONC). X Region: Valdivia, Chihuio, Hito de Portezuelo Ipela, Goday 119 (CONC); Palena, Las Escalas, Futaleufu, Hildebrand-Vogel 31 (CONC); Rio Chico, Futaleufu, Hildebrand-Vogel 45 (CONC). XI Region: Aysen, Reserva Forestal Mano Negra, Schlegel 7194 (CONC); Reserva Forestal Mano Negra, Schlegel 7187 (CONC); Prov. Coihaique, Sector Lago Palena, Godoy, Hildebrand-Vogel & Vdgel 3 (CONC); Parque Nacional Trapananda, Schlegel 8070 (CONC); General Carrera, Estero Cofre, Vogel 540 (CONC); Estero Cofre, Vogel 519 (CONC); Rio Ibanez, Vogel 5 (CONC); Capitan Prat, Villa O'Higgins, Vogel s.n. (CONC). XII Region: Magallanes, Punta Arenas, Minas de carbon, Ricardi & Matthei 335 (CONC); Puerto Williams, Isla Navarino, Cerro Bandera, Schlegel 8122 (CONC). 2. Trisetum flavescens (L.) P. Beauv., Ess. Agrostogr. 88, 153, t. 18, f.l. 1812. Basionym: Avena flavescens L., Sp. pi. 80. 1753. Trisetaria flavescens (L.) Baumg., Enum. Stirp. Transsilv. 3: 263. 1816. Rebentischia flavescens (L.) Opiz, Volume 92, Number 4 2005 Finot et al. Trisetum in South America 541 Lotos 4: 104. 1854, as synonym of Trisetum flavescens (L.) P. Beauv. TYPE: Herb. A. Van Royen no. 913.7^58 (lectotype, LINN-97.14, designated by Cope in Cafferty et al., Taxon 49(2): 247. 2000, not seen). guna Mucubaji, toward Laguna Negra, 3625• 3655 m, 21 July 1944, J. A. Steyermarh 57482 (holotype, US-1911640!; isotypes, M03846028!, MO-3873805 fragm. ex VEN!; VEN not seen, F-1216139!). Perennial, caespitose; culms (20•)80•110 cm tall; nodes 2 to 5, glabrous. Leaf sheaths shorter than the internodes, glabrous or sparsely pilose below; ligule 0.5•2 mm long, truncate, ciliate, the dorsal surface glabrous; blades (3•)10•16 X 0.2• 0.4 cm, flat, glabrous abaxially, scabrous and sparsely pilose adaxially. Panicles 5•18 X 2•8 cm, dense, lax, open or narrow, gold-yellow, shining. Spikelets 5•7(•8.5) mm, 2- to 4-flowered; rachilla ca. 1.2 mm long, pilose, with trichomes up to 1 mm long; glumes shorter than the florets, unequal; keel scabrous; first glume 2•4 X 0.1•0.2 mm, linearlanceolate, narrow, 1-nerved, subulate, usually 1/2 as long as the second glume; second glume 4•6.6 X 1 mm, oval-lanceolate, 3-nerved; lemmas 4•6 mm long, glabrous, awned, somewhat scabrous toward the apex; apex 2-dentate to 2-aristulate; awn 5•9 mm long, borne on upper 1/3 or 1/4, geniculate and twisted toward the base; callus pubescent, the trichomes ca. 0.5 mm long; paleas a little shorter than the lemma, hyaline, 2-nerved, the nerves scabrous; apex 2-dentate to 2-setulate; lodicules bilobed at the apex; anthers 2•3 mm long; ovary glabrous. Caryopses 2•3 mm long, glabrous; endosperm liquid. Perennial, with short rhizomes; culms glabrous, 35•80 cm tall; nodes glabrous, basal. Leaf sheaths sometimes with auricles, lower sheaths covered with short retrorse trichomes, the upper sheaths glabrous to puberulous; ligule 2•4 mm long, hyaline, oval to triangular, denticulate on the margins, the dorsal surface glabrous; blades 7•20 X 0.1•0.6 cm, flat; lower blades on culm densely pubescent, upper blades glabrous. Panicles 10•19 X 1•1.5 cm, lax, contracted, not spiciform, linear, purple; rachis glabrous. Spikelets 7•7.5 mm long, 2-flowered; pedicels 0.5•5 mm long, glabrous; rachilla 1• 1.8 mm long, covered with trichomes 1•1.5 mm long; glumes subequal, the first glume a little shorter than the second glume, membranous; margins hyaline; keel scabrous near apex; apex acute; first glume 5.5•6.4 X 0.6•1.2 mm, oval-lanceolate, attenuate, 1-nerved; second glume 6.5•7 X 1•1.6 mm, oval, 3-nerved, almost equaling the spikelet in length; florets 6•6.7 X ca. 1 mm; upper floret shorter; lemmas glabrous, awned dorsally; apex bidentate, the teeth aristulate with projection of the intermediate nerves; awn 8•11 mm long, borne on the upper 1/3, at 2•2.5 mm from the apex, twisted at the base, curved or weakly geniculate, scabrous; callus obtuse, with trichomes 0.5•0.8 mm long; paleas ca. 4.5 mm long, shorter than the lemma, hyaline, 2-nerved, the nerves scabrous; anthers 1.2• 1.5 mm long; lodicules 0.7•0.8 mm long, obovate, with 3 or 4 short apical teeth; ovary glabrous, the styles well apart. Caryopses not seen. Chromosome number. In = 24, 28 (Bolkhovskikh et al., 1969; Tsvelev, 1983; Dixon, 1995; Frey, 1992). Distribution and habitat. A European species that was introduced from Europe in North and South America (Frey, 1992). In South America it occurs in the Chilean and Argentinean Patagonia and is an important forage plant (Nicora, 1978). Zuloaga et al. (1994) cited this species for Neuquen, Argentina. Phenology. Flowering in December. Illustrations. Hitchcock and Chase (1950: 291, fig. 393); Nicora (1978: 242, fig. 157 A-C). Additional specimens studied. ARGENTINA. Neuquen: Peninsula Panguinal, costa N Lago Nahuel Huapi, Diem 463 (BAA); Nahuel Huapi, cultivated, Gallinal, Aragone, Bergalli, Campal & Rosengurtt 5594 (US). GERMANY. Magdeburg, 23 lune 1932, Schwing s.n. (CONC). U.S.A. Washington: Walla Walla, 31 May 1900, Lechenby s.n. (US). 3. Trisetum foliosum Swallen, Contr. U.S Natl. Herb. 29(6): 256. 1948. TYPE: Venezuela. Merida: rocky slopes along stream above La- Distribution and habitat. Trisetum foliosum is endemic to Venezuela. It has been collected in paramos on rocky slopes and near streams and lakes, between 3500 and 3655 m. Phenology. Flowering between November and July. Comments. Trisetum foliosum resembles T. flavescens and T. cernuum in having lax panicles and spikelets with glumes shorter than the florets. It differs from T. cernuum in having subequal glumes, the first glume a little shorter than the second glume, the first glume never reduced, and the ovary is glabrous (glumes very unequal, the first glume conspicuously shorter than the second glume, the first glume sometimes reduced, and the ovary is hairy in T. cernuum). Trisetum foliosum differs from T. flavescens in having more contracted (1•1.5 cm wide) and purple panicles and longer ligules (2-4 542 Annals of the Missouri Botanical Garden mm long) versus open (2•8 cm wide) and gold-yellow panicles, and shorter ligules (0.5•2 mm long) in T. fla tavescens. and geniculate; callus obtuse, with short trichomes, the trichomes ca. 0.2•0.3 mm long; paleas 3.2•4.6 mm long, slightly shorter than the lemma, hyaline, 2-keeled, the keels scabrous on the upper half, 2dentate to 2-awned at the apex; anthers (0.8•)1.2• 1.8 mm long; lodicules ca. 0.7 mm long, bilobed at the apex, the lobes acute; ovary glabrous. Caryopses 2.2•3 mm long; endosperm soft. Additional specimen studied. VENEZUELA. Merida: pass on the Merida•Barinas Hwy., paramo above Laguna Grande and Universidad de Los Andes Experimental Station, Davidse 3226 (US). 4. Trisetum irazuense (Kuntze) Hitchc, Proc. Biol. Soc. Wash. 40: 82. 1927. Basionym: Calamagrostis irazuensis Kuntze, Revis. Gen. PL 2: 763. 1891. TYPE: Costa Rica. Volcan Irazu, Waldregion, 3000 m, 24 June 1874, C. E. 0. Kuntze 2334 (holotype, NY-346300 ex Herb. Kuntze!). Trisetum scabriflorum Hitchc., Contr. U.S. Natl. Herb. 24: 358. 1927. TYPE: Colombia. Cauca: collected below Pitayo, Rio Palo Basin, Tierra Adentro, 2400 m, Feb. 1906, H. Pittier 1435 (holotype, US-531631!). Trisetum fournieranum Hitchc, Contr. U.S. Natl. Herb. 17: 326. 1913, new name for Trisetum gracile E. Fourn., Mexic. PI. 2: 108. 1886, nom. Meg. TYPE: "San Luis de Potosi," M. Virlet dAoust 1382 (lectotype, P!, designated by Finot et al., Ann. Missouri Bot. Card. 91: 13. 2004; isotype, US-726971b fragment ex P!). Perennial; caespitose; culms 75•100 cm tall, glabrous, up to 2 mm diam. on the lower internodes; nodes 3, glabrous. Leaf sheaths shorter or slightly longer than the internodes, glabrous or pubescent; basal sheaths pubescent; ligule 1•^ mm long, membranous, pubescent to densely pubescent, the apex truncate, dentate or ciliate; blades 20•30 cm X 2.5•6 mm, flat, glabrous or pubescent; upper blades 5•10 cm long. Panicles (7•)3•25 X (1•)2•5 cm, lax, narrow, yellow-green to deep green and purple; rachis glabrous to sparsely pubescent; the branches appressed and ascending. Spikelets 4•7(•9) mm long, 2- or 3-flowered; pedicels 2•7 mm long, glabrous to sparsely pubescent, sometimes scabrous; rachilla 1•1.7 mm long, covered with stiff trichomes, the trichomes 0.5•1 mm long; glumes very unequal, shorter than the florets; the keel scabrous on the upper half, the margins hyaline; first glume 2.5•5 mm long, narrow, linear, 1nerved, acute at the apex, half as long as the spikelet; second glume 3.5•6.5 mm long, lanceolate to ovate-lanceolate, abruptly attenuate, 3-nerved, 3/4 as long as the spikelet; lemmas 4.3•6 mm long, 5nerved, the nerves inconspicuous toward 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 1/3 or 1/4, straight or twisted Chromosome number. In = 28, 42 (HernandezTorres & Koch, 1988). Anatomy and micromorphology. HernandezTorres & Engleman (1995); Finot et al. (2004). Distribution and habitat. Trisetum irazuense ranges from Mexico to Peru (Calderon de Rzedowski & Rzedowski, 2001; Finot et al., 2004). In South America it is found in Colombia, Venezuela, Ecuador, and Peru between 1500 and 3800 m. Phenology. Flowering between November and August. Illustrations. Pohl, W. (1980: 578, fig. 217 C). Comments. Trisetum irazuense resembles T. cernuum in having lax, open panicles bearing long pedicellate spikelets with dissimilar glumes shorter than the florets. Anatomically both species have very similar characteristics in anatomical transverse section, and abaxial and adaxial epidermes. However, these two species are easily separated. Trisetum irazuense has strongly scabrous lemmas (relatively glabrous in T. cernuum) and a glabrous ovary (hairy in T. cernuum) (see additional comments under T. cernuum). Additional specimens studied. COLOMBIA. Bogota: Pena, Lindig 1862 (P). Caldas: Laguneta, Salento, von Sneidern 3027 (S). Santander: Minas San luan 5 km above California, Robinson & Beltran 3041 (US). Departamento del Valle: Cordillera Central, vertiente occidental, Hoya del no Bugalagrande, Barragan, cerro de La Laguna, Cuatrecasas 20847 (F); Cordillera Central, vertiente occidental, Hoya del no Bugalagrande, Loma de Barragan, desde La Parrilla a La Machuca, Cuatrecasas 20680 (F). ECUADOR. Carchi: Hacienda La Esperanza, sector El Voladero, paramo de El Angel, Ddvalos 20 (US); 22 km SW of Tulcan on road to El Angel, Peterson et al. 9146 (US); Valle de Maldonado, km 53 on the road Tulcan• Maldonado, Holm-Nielsen et al. 5553 (S); Las Penas between La Rinconada and San Gabriel, Asplund 7215 (S); above el Pun toward Tulcan, Asplund 16868 (S). Picliincha: below San luan, Asplund 16093 (S); vicinity of Quito, Asplund 6156 (S); Mt. Corazon, paramo, Asplund 9686 (S); Nono, Asplund 7441 (P, S); Reserva Geobotanica Pasochoa, Laegaard 101414 (QCA); Pululagua, Asplund 6733 (US). Riobamba: Hacienda Toldo, Oct. 1891, Sodiro s.n. (P). Azuay/Morona: near the pass on road Sigsig•Gualaquiza, Laegaard et al. 103031 (QCA). Bolivar: La Magdalena, Asplund 8280 (P, S). Tungurahua: vicinity of Patate, Hacienda Leito, Asplund 7978 (P). PERU. Piura: Prov. Huancabamba, 23 km E of Sondor, on road toward Tabacones, Peterson & Refulio-Rodriguez 15135 (US). VENEZUELA. Merida: entre Timotes y Chachopo, Bur- Volume 92, Number 4 2005 Finot et al. Trisetum in South America kart & Tamayo 16733 (BAA). Monagas: Cerro Negro, above La Sabana de las Piedras, NW of Caripe, Steyermark 62083 (F). Sucre: Cerro Turumuquire, north-facing slopes between La Trinidad and zone of cloud forest, Steyermark 62488 (F). in Region XII, Prov. de Ultima Esperanza, between 50°S and 51°S, and between 500 and 900 m. Phenology. Flowering between January and February. Illustrations. Rugolo de Agrasar and Nicora (1988: 469, fig. 2A-K). Comments. Trisetum ambiguum is closely related to T spicatum var. cumingii and differs from the latter by having 1-flowered spikelets, isomorphic glumes, the awn several times curved, and the ovary and caryopsis with a few trichomes. Trisetum spicatum var. cumingii has 2- or 3-flowered spikelets, unequal glumes, a geniculate awn, and a glabrous ovary and caryopsis. Trisetum subg. Trisetum sect. Trisaetera Asch. & Graebn., Syn. Mitteleur. Fl. 2: 270. 1895. TYPE: Trisetum spicatum (L.) K. Richt. 5. Trisetum ambiguum Rugolo de Agrasar & Nicora, Bol. Soc. Argent. Bot. 25: 468. 1988. TYPE: Argentina. Prov. Santa Cruz: Dpto. Giier Aike, Ea. Sofia, Secc. Cuadrado, 5 km al S de Est. Punta del Monte, 350 m, 51°41'S, 71°18'W, 12 Feb. 1978, O. Boelcke, D. Moore & F. Roig 3119 (holotype, BAB not seen; iso- type, SI!). Perennial, with short rhizomes; culms 12•34 cm tall, with short, retrorse trichomes below the inflorescence. Leaf sheaths glabrous or pubescent; ligule 1•2.5 mm long, denticulate, laciniate, the dorsal surface glabrous; blades 40•50 X 1.5•2 mm, short, flat to conduplicate, scabrous on the margins, glabrous to pilose abaxially, scabrous-pubescent to pilose adaxially. Panicles 4•7.5 X 1•1.5 cm, spiciform, dense; rachis pilose. Spikelets ca. 6 mm long, 1-flowered; pedicels scabrous; rachilla ca. 1.5 mm long, pilose with trichomes 1.5 mm long; glumes longer than the floret isomorphic; first glume 5•5.5 X ca. 0.5 mm, equal or slightly shorter than the floret, and slightly shorter and narrower than the second glume, lanceolate to oval-lanceolate, equaling or slightly shorter than the florets, 1nerved; second glume 5•5.5 X ca. 0.8 mm, slightly longer or equaling the floret, lanceolate to oval-lanceolate, 3-nerved; florets 4•5 mm long; lemmas glabrous, scabrous toward the apex, 5-nerved, the nerves conspicuous toward the apex, awned dorsally; apex hyaline with the intermediate and marginal nerves prolonged as 4 short awns; awn 4•5 mm long, borne on the upper 1/3, strongly twisted and curved; callus obtuse, with trichomes ca. 0.5 mm long; paleas 3•3.5 mm long, shorter than the lemma, 2-nerved, the nerves scabrous toward the apex; flowers cleistogamous; anthers 0.5•0.8 mm long; lodicules ca. 0.8 mm long, minutely bilobed near the apex; ovary with 1 to several curved trichomes at the apex. Caryopses ca. 2.5 mm long, pubescent at the apex; endosperm dry. Distribution and habitat. Endemic to southern Argentina and Chile (Zuloaga et al., 1994; Finot, 2002). In Argentina it is found in Santa Cruz and Tierra del Fuego, between 51°S and 53°55'S (Rugolo de Agrasar & Nicora, 1988). In Chile it grows 543 Additional specimens studied. ARGENTINA. Santa Cruz: Giier Aike, Valle Superior Rio Turbio, Ambrosetti & Me'ndez 29875 (MERL); Valle Superior Rio Turbio, Ambrosetti & Mendez 29883 (MERL); Estancia La Carlota, seccion San Ellas, Roig et al. 103 (SI). Tierra del Fuego: Norte de Rio Grande, Ea. Maria Behety, Soriano 4847 (BAA, paratype); Castillo, 30 Jan. 1942, Castellanos s.n. (BA); Rio Grande, 1 Mar. 1917, Bonarelli s.n. (SI); Cerro Mesa, 31 Jan. 1942, Castellanos s.n. (BA). CHILE. XII Region: Prov. de Ultima Esperanza, Sierra de los Baguales, La Cumbre, Campo de La Tropilla, Landero 790 (CONC); Sierra de los Baguales, Cerro Santa Lucia, 15 Jan. 1985, Arroyo s.n. (CONC 85160); Sierra del Tore, Arroyo et al. 9276 (CONC); Sierra del Toro, 10 Feb. 1992, Arroyo et al. s.n. (CONC 92112). 6. Trisetum andinum Benth., PI. Hartw. 261. 1847. Trisetum spicatum var. andinum (Benth.) Louis-Marie, Rhodora 30: 239. 1929. Trisetum spicatum (L.) K. Richt. subsp. andinum (Benth.) Hulten, Svensk Bot. Tidskr. 53: 224. 1959. TYPE: Ecuador. Hacienda de Antisana, C. T. Hartweg 1449 (holotype, K not seen; isotypes, P!, BAA-3345!, US fragm. ex PSTEUD!). Perennial, caespitose, velvety, sometimes with short rhizomes; culms (9•)20•42 cm tall, erect, robust, up to 4 mm diam. near the base, pubescent almost all its length, densely hairy below the panicle; nodes 1, basal. Lower leaf sheaths ca. 20 cm long, lax, pilose; upper sheaths 5•10 cm long, pilose; ligule 0.5•2 mm long, truncate, ciliate on the margin, dorsal surface densely pilose; blades 3•15 cm X 2•3.5 mm, flat, pilose abaxially, sparsely pilose adaxially; margins of lower blades usually involute. Panicles 3•6 X 1•2.5 cm, spiciform, very dense, ovate, no more than 3 times as long as wide, included or exerted in the upper sheath, tinged with gold and purple, rounded at the apex; rachis pubescent; spikelets 5•7 mm long, 2- or 3-flowered; rachilla 0.6•0.8 mm long, with a few stiff trichomes, the trichomes as long as the rachilla; 544 Annals of the Missouri Botanical Garden glumes equaling or longer than the florets, sometimes only the first glume longer than the florets, unequaf; first glume 3.5•6 X 0.5•0.6 mm, shorter and narrower than the second glume, linear-lanceolate, attenuate, 1-nerved; second glume 4•7 X 0.8•1 mm, oval, 3-nerved, rarely shorter than the florets; florets 5•5.5 mm long; lemma scabrous or covered with very short trichomes giving a velvety texture, dorsally awned, purple and green, the apex 2-aristulate with the projection of the intermediate nerves, the mucros 0.4•0.5 mm long; awns 3.5•4.5 mm long, inserted on upper 1/3, usually not geniculate nor twisted but diversely curved, scabrous; callus obtuse, with few short trichomes, the trichomes ca. 0.2 mm long; paleas 3.7•6 mm long, shorter than the lemma, hyaline, 2-nerved, the nerves scabrous; apex 2-dentate, sometimes shortmucronate; anthers 0.7•1 mm long; lodicules ca. 0.8 mm long, hyaline, 2-lobulate at the apex; ovary glabrous. Caryopses 2.2•2.5 X ca. 0.5 mm, glabrous. cano, Asplund 7499 (S). Ibambura: NE side of Cayambe Mountain, 10 Dec. 1961, Cazalet & Pennington 5740 (US); W side of Mount Cayambe, in paramos, Little & Paredes 68321'2 (US). Pichincha: Pichincha, Andre 3907 (US); Summit of Pichincha, Jameson s.n. (US-868479); Mt. Pichincha, near Quito, Hitchcock 21059 (US); 36 km E of El Refugio, on the SW slope of Volcan Cayambe, Peterson, Judziewicz & King 9075 (US); Faldas SE Volcan GuaguaPichincha, Nowak & Marcillo 79 (QCA); Guagua Pichincha, E slopes of the volcano, Sklenar & Kosteckova 812 (QCA); Guagua Pichincha, Asplund 7401 (S, US); W parts of Rucu Pichincha massif, summit and upper W slopes of Padre Encantado, Molau & Eriksen 3297 (QCA); Guagua Pichincha, Hading 4537 (S); Lloa-Guagua Pichincha km 10, Laegaard, Romoleroux & Leon 102731 (QCA); sommet du Pichincha, Oct. 1856, /. Remy s.n., (P); Pichincha, Benoist 2389, 4389 (P); monte Rucu Pichincha, Holmgren 553 (S). Pichincha/Napo: Antisana, Ehrenburg 137 (QCA); Volcan Antisana, between Campamento IMAP and Laguna Micacocha, Laegaard 101604 (QCA). Distribution and habitat. Endemic to Ecuador, Trisetum andinum is found between 3900 and 4700 m, in paramo and superparamo vegetation, on volcanic soil. Phenology. Flowering between December and September. Comments. Trisetum andinum resembles T. spicatum. It has been treated as a synonym of T. spicatum by j0rgensen and Ulloa (1994), Renvoize (1998), and j0rgensen and Leon-Yanez (1999). Hitchcock (1927) accepted T. andinum as a good species, but it was later treated by Louis-Marie (1928, 1829) as a variety and by Hulten (1959) as a subspecies of T. spicatum. The most remarkable differences between T. andinum and T. spicatum are the shape of the panicles and the hairiness of the plant. Trisetum andinum is totally covered with a dense indumentum (T. spicatum is glabrous except below the panicle). Trisetum andinum possesses oval, dark purple panicles, usually no more than 2 or 3 times longer than wide (T. spicatum possesses panicles green or purple, narrow, more than 3 times longer than wide). Trisetum andinum possesses glumes as long as the florets or the second glume longer than the florets (glumes shorter than the florets in T spicatum). The isotype of T. andinum at P has both glumes longer than the florets (first glume 6 X 0.8 mm, second glume 7 X 0.8 mm). Additional specimens studied. ECUADOR. Cotopaxi: P.N. Cotopaxi, Ehrenburg 49 (QCA); Volcan Cotopaxi, Sklenar & Kosteckova 80•12 (QCA); Paramos del Cotopaxi, Roig 12356 (MERL); Cotopaxi, SW slope of the vol- 7a. Trisetum barbinode Trin. var. barbinode, Linnaea 10(3): 300. 1836. TYPE: Chile Austral. Andes de Antuco, 1828, E. Poeppig s.n. (holotype, LE-TRIN-1886.01!; isotypes, US81770 ex W!, US-868486 fragm. ex LETRIN!, BAA-3351 fragm. ex LE-TRIN!, SGO73101 photo ex LE-TRIN!, W not seen). Perennial; culms (18•)25•45(•55) cm tall, sometimes with short rhizomes, pilose or sericeous below the panicle, the trichomes first antrorse, then retrorse; nodes 1 or 2, basal, pubescent. Leaf sheaths glabrous; ligule 0.5•2 mm long, truncate, dentateciliate, dorsally glabrous; blades (2•)10•15 cm X 2•3(•5) mm, flat, soft, glabrous, margins and adaxial surface scabrous. Panicles 4.5•15 X 0.8• 2.5(•3) cm, pale green, contracted, dense, linearlanceolate to oval; rachis pubescent, the trichomes up to 1.5 mm long; spikelets 6•8(•11) mm long, 2or 3-flowered; pedicels up to 5 mm long, pubescent; rachilla 1•2 mm long, pubescent, the trichomes up to 3 mm long; glumes unequal, with hyaline margins, the apex shortly aristulate or acute, the keel scabrous on the upper half, both glumes or only the second glume longer than the florets; first glume (5.5-)6.5-8.5 X 0.5-0.9 mm, slightly shorter and narrower than the second glume, linear-lanceolate, 1- or 3-nerved, usually equaling the first floret, sometimes longer; second glume 6.5•10 X 0.7•1.2 mm, lanceolate, 3-nerved; florets (5•)6.5•7.5 X (0.5•)0.8•1 mm; lemmas pubescent, dorsally awned, the apex biaristulate with apical awns 0.5• 1 mm long; awn (2•)6.5•10(•14) mm long, curved or geniculate, not twisted; callus obtuse, pubescent, the trichomes 1.5•3 mm long; paleas (4•)5•6.5 mm long, shorter than the first lemma, equal or longer than the lemma in the second or third floret, hyaline, 2-nerved, the nerves scabrous, the apex bi- Volume 92, Number 4 2005 Finot et al. Trisetum in South America dentate, the teeth 2-setulate; anthers (0.7•)1•1.6 mm long; lodicules 0.7•1 mm long, 2- or 3-lobate at the apex; ovary glabrous or with a few short trichomes at the apex. Caryopses 2.5•3.5 mm long, glabrous or with few trichomes at the apex; endosperm liquid. type fragment at US it is not possible to distinguish T. sclerophyllum from T. barbinode Trin. The varietal status for T sclerophyllum was also suggested by Parodi since there is an annotation label on the type fragment deposited at BAA with his script. This, also, was never published. Distribution and habitat. Endemic from central-southern Chile and Argentina, it is distributed along the Cordillera de los Andes, between 36°S and 41°S. In Argentina, T. barbinode var. barbinode grows in Neuquen, Rio Negro, and Chubut, and in Chile between the Regions VII and IX, usually between 550 and 3500 m, on volcanic soils. Phenology. Flowering between January and March. Illustrations. Nicora (1978: 256, fig. 165A-C). Comments. Morphologically, Trisetum barbinode var. barbinode is characterized by having contracted, pale green panicles with large spikelets 6• 8(•11) mm long, hairy lemmas, and 3-nerved glumes longer than the florets. Trisetum barbinode resembles other taxa with hairy lemmas, such as T. sclerophyllum and T. hirtiflorum. Due to the variation in the shape, color, and size of the panicle and the length of the spikelets and glumes, it is often extremely difficult to distinguish among these three species. Trisetum hirtiflorum resembles T. barbinode s. str., and the only difference between these two species is the presence of a few short hairs at the apex of the ovary and caryopsis in T hirtiflorum. According to Nicora (1978), T hirtiflorum has shorter and narrower panicles and more rigid leaves than T. barbinode. However, these characters are so variable that recognition of these two species is nearly impossible. Louis-Marie (1929) first treated T. hirtiflorum as a variety of T. barbinode. The close relationship between Trisetum sclerophyllum and T. barbinode was first observed by Louis-Marie, as it is clear from the annotation he made on the type deposited at US: "7] barbinode var. sclerophyllum" signed "Lalonde." However, Louis-Marie (1928, 1929) did not include this combination or the Hackel binomial in his treatment of genus Trisetum. This type fragment at US from W contains only a fragment of the panicle. The spikelets are 2-flowered; the glumes are nearly equal and longer than the florets, the first glume 6•8 X 0.6• 0.7 mm, the second glume 7•8 X 0.7•1 mm, and both are 3-nerved, with scabrous keel, hyaline margins, and hyaline, aristulate apex. The lower floret is ca. 5 mm long and the lemmas are pubescent, with a bi-aristulate apex, with apical awns up to 1 mm long. The dorsal awn (6•8 mm long) is borne on the upper 1/3 of the lemma. On the basis of this Additional specimens studied. ARGENTINA. Neuquen: Catan Lil, Sierra del Chachil, Riigolo de Agrasar 451 (BAA); Loncopue, Cajdn Chenque Pehuen, Riigolo de Agrasar & Agrasar 148 (BAA); Los Lagos,Villa La Angostura, cerro Belvedere, Riigolo de Agrasar 1234-1 (SI); P.N. Lamn, Volcan Huanquihue, al W del Lago Currhue. Eskuche & Klein 1520-7 (CORD); a 5 km de Las Ovejas, camino a las lagunas Epu-Lauquen, Boelcke 10764 (CONC ex BAA); Bafios Calientes, Rio Valvarco, Boelcke et al. 114521/2 (BAA); entre Lagunas Epu-Lauquen y Cajdn Lumavia, Ragonese 234 (BA); Lagunas Epu-Lauquen, Boelcke et al. 10927 (CONC ex SI). Rio Negro: P.N. Nahuel Huapi, Cuenca Rio Manso Superior, camino a Ventisqueros del Tronador, Diehl & Bravo 10843 (BA). CHILE. VII Region: Talca, Laguna del Maule, Schlegel 3475 (CONC); Laguna del Maule, Schlegel 3537 (CONC). VIII Region: Nuble, Termas de Chilian, Jaffuel 1806-b (SCO), 9 Jan. 1949, Pfister s.n. (CONC); Bafios de Chilian. Jan. 1877, F. Philippi s.n. (SGO); Bafios de Chilian, Philippi 218 (SGO); Termas de Chilian, Faldeos del Volcan Chilian, Ricardi 5604 (CONC); Termas de Chilian, Philippi 229 (SGO); Termas de Chilian, zona de las fumarolas, Finot & Baeza 2069 (CONC); entre Valle Hermoso y Termas de Chilian, Finot & Baeza 2070 (CONC); Refugio Club Andino de Chilian, Shangri-La, Garaventa 4681 (CONC); Bio-Bio, Laguna del Laja, Fabris & Crisci 7608 (LP); Lago Laja, 2 Feb. 1968, Zoellner s.n. (ZOELLNER): Laguna del Laja, Ricardi & Marticorena 5812/1973 (CONC), Burkart 27451 (CONC ex SI); faldeos del Volcan Antuco, frente a la laguna del Laja, Ricardi & Marticorena 5723/1884 (CONC); Las Lagartijas, Finot & Baeza 2073 (CONC). IX Region: Malleco, Lonquimay, Estepa cerca Laguna Icalma, 10 Jan. 1947, Pfister s.n. (CONC); Curacautin, Paso Las Raices, 14 Feb. 1992, Zoellner s.n. (ZOELLNER); Termas de Rio Blanco, Pfister s.n. (CONC); Icalma, cerro del lado sur, Pfister s.n. (CONC); Cautin, Volcan Llaima, Refugio Cautin, Montero 4497, 4487 (CONC); Volcan Llaima, Gunckel 12426 (CONC), Sparre 4873 (S); Villarrica, Tromen, Limite Chileno-Argentino, Ricardi & Matthei 3, 15 (CONC); Curarrehue, 29 Dec. 1946, Canulafs.n. (CONC). 7b. 545 Trisetum barbinode var. hirtiflorum (Hack.) Louis-Marie, Rhodora 30: 240. 1929. Basionym: Trisetum hirtiflorum Hack., Repert. Spec. Nov. Regni Veg. 10 (243-247): 169. J9JJ. TYPE: Chile. C. Reiche s.n. (holotype, W not seen; isotypes, BAA!, SGO fragm. & photo ex W!, US fragm. ex W!). Panicles 4•5 cm long, gold-purple, spiciform, dense. Leaf blades soft. Second glume 5•8 mm long; ovary with short trichomes on the apex. Distribution and habitat. Southern Chile (VIII Region) and Argentina (Neuquen) between 500 and 2000 m (Nicora, 1978). 546 Annals of the Missouri Botanical Garden Phenology. Flowering in January. Additional specimen studied. CHILE. VIII Region: Prov. Nuble, Termas de Chilian, por la loma de las fumarolas, 17 Jan. 1945, Pfister s.n. (CONC, SGO). dulata (Trin.) Griseb., Abh. Konigl. Ges. Wiss. Gottingen 24: 292. J879. TYPE: Chile. Andes Chile boreal, E. F. Poeppig s.n. (holotype, LETRIN-J887.0J!; isotype, BAA-3364 fragm. ex LE-TRIN!). 7c. Trisetum barbinode var. sclerophyllum (Hack.) Finot, comb. nov. Basionym: Trisetum sclerophyllum Hack., Anales Mus. Nac. Hist. Nat. Buenos Aires 2J: J08. J9JJ. TYPE: Argentina. Chubut: Dpto. Languineo, Region del rfo Corcovado, 20 Jan. J902, Illin 148 (holotype, W ex Stuckert Herb. Arg. J799J not seen; isotypes, BAA-34J4!, US-9J365 fragm. ex W!). Culms J8•30 cm tall. Leaf blades stiff. Panicles (2•)3•5(•8) cm long, gold-purple, spiciform, dense. Glumes subequal in width; second glume 5•8 mm long; ovary glabrous. Distribution and habitat. Endemic to Argenti- na, T. barbinode var. sclerophyllum occurs along the Cordillera de Los Andes in Mendoza, Neuquen, Rio Negro, and Chubut, between 32°S and 42°S latitudes and between JJ70 and 3500 m. Phenology. Flowering between January and March. Additional specimens studied. ARGENTINA. Mendoza: Las Heras, Faldeo SSW morrena cerro Tolosa, Boelcke et al. 9778 (MERL, CONC); San Rafael, Cerro Volcan Overo, Roig 1 (MERL); alto valle de Calmuco, Burhart et al. 13930 (BAA ex SI); alto valle del Atuel, Paci 15731 (BAA, MERL); Laguna Atuel, Ruiz Leal 16886 (MERL); Tupungato, Las Tres Quebradas, Ruiz Leal 3616 (MERL). Neuqen: Sierra de Cochico, Boelcke et al. 14083 (BAA, CONC ex SI); Norquin, entre Las Maquinitas y Copahue, Calderon & Riigolo 57 (BAA); Copahue, Calderon & Riigolo 72 (BAA); Pino Hachado, Feb. 1920, Hauman s.n. (BA), Parodi 2700 (BAA); Paso Pino Hachado, en el hito, Nicora 7432 (BAA); San Martin de Los Andes, P.N. Lanin, arroyo Rucu-Leufu, Correa et al. 5647 (CONC ex BAB); P.N. Lanin, Cerro Chapelco, 12 Feb. 1961, Leon & Calderon s.n. (BAA); Cerro Chapelco, Schajowski 86, 133, 134, 178 (BA), Cabrera & Crisci 19145 (LP), Cabrera et al. 23024 (LP); Lago Lacar, Cerro Malo, Riigolo de Agrasar 315, 318 (BAA); Cerro Repollo, Estancia Meliquina, Riigolo & Agrasar 570 (BAA); Los Lagos, Villa La Angostura, cerro Belvedere, Riigolo de Agrasar 1234•2 (SI); Pto. Manzano, en la cima del cerro O'Connor, Diem 3229 (BAA); P.N. Nahuel Huapi, Filo refugio Cerro Colorado a Cerro, Boelcke & Correa 6959 (BAA); mallines en ladera Cerro, Boelcke & Correa 6964 (BAA). Rio Negro: Cerro de Las Hormigas, Hosseus 559, 560 (CORD); P.N. Nahuel Huapi, Cerro Catedral, Cabrera 19754 (LP), Parodi 15321 (BAA), PerezMoreau s.n., 1949 (BA). Prov. Chubut: Dpto. Futaleufu, Esquel, La Hoya, Cabrera et al. 23170 (LP); P.N. Los Alerces, Lago Futalaufquen, Soriano 4352 (BAA). 8a. Trisetum caudulatum Trin. var. caudulaIIIIII. Linnaea J0(3): 300. J836. Koeleria cau- Trisetum chromostachyum E. Desv., Fl. Chil. 6: 350. 1854. TYPE: Chile. Santiago, in arvis, Jan. 1829, C. Gay s.n. (holotype, P!; isotypes, CONC-148153 fragm. ex P!, SGO fragm. ex P!, US fragm. ex P-DESV-156L US fragm. ex P-GAY!). Trisetum heteronymum Steud., Syn. PL Glumac. 1: 229. 1854. TYPE: Chile. "Rancagua, in pascuis et ad fossas," Oct. 1828, C. L. G. Bertero 53 (published as Bertero 83) (holotype, P-STEUD-437!; isotypes, SGO fragm. ex P!, US fragm. ex P!; US fragm. ex P!). Trisetum variabile E. Desv., Fl. Chil. 6: 351. 1854. Trisetum variabile var. (alpha) flavescens E. Desv., FL Chil. 6: 351. 1854. TYPE: (Chile) Valparaiso, 1829, C. L. G. Bertero 998 (lectotype, designated here, P!; isotypes, BAA-3422 fragm. ex P!, SGO fragm. ex P!, US fragm. ex P!, US fragm. ex P-STEUD-440!). Trisetum variabile var. virescens E. Desv., Fl. Chil. 6: 351. 1854. TYPE: Chile. Prov. Valdivia, In herbosis Guanegue, Feb. 1889, C. Gay s.n. (lectotype, designated here, P!; isotypes, BAA-3423 fragm. ex P!, CONC fragm. ex P!; SGO fragm. ex P!, US fragm. ex P!). Trisetum malacophyllum Steud., Syn. PL Glumac. 1: 229. 1854. non Phil., Anales Univ. Chile 48: 566. 1873. TYPE: (Chile) Valparaiso, "Festuca nr. 997 Hbr. Bertero, In sylvaticis calidis Collinum loco dicto La Laguna," Oct. 1829, C. L. G. Bertero 991 (lectotype, designated here, P!; isotypes, GH not seen, SGO fragm. ex P!). Trisetum splendidulum Steud., Syn. PL Glumac. 1: 229. 1854. TYPE: Chile. "Festuca" C. L. G. Bertero 996 (holotype, P-STEUD-441!; isotypes, BAA-3416 fragm. ex P!, CONC fragm. ex P!, SGO fragm. ex P!, US fragm. ex P!). Trisetum chiloense Phil., Linnaea 29: 93. 1858. Trisetum variabile E. Desv. var. chiloense (Phil.) Louis-Marie, Rhodora 30: 240. 1928. TYPE: Chile, "in pascuis insulae Chiloe, ad Castro" C. Gay 147 (holotype. SGO-PHIL-215!; isotypes, BAA-3371 ex SGO!, US fragm. ex SGO-PHIL-215 & photo!). Trisetum monticola Phil., Linnaea 33: 291. 1864. TYPE: Chile. Prope Santiago, in andibus, Nov. 1861, R. A. Philippi s.n. (holotype, SGO-PHIL-227!; isotypes. SGO-37049!; US fragm. & photo ex SGO-PHIL227!). Trisetum ochrostachyum Phil., Linnaea 33: 290. 1864. TYPE: Chile. "E Valdivia attuli," R. A. Philippi s.n. (holotype, SGO-PHIL-220!; isotype, SGO-37062!). Trisetum vidalii Phil., Anales Univ. Chile 94: 27. 1896. Trisetum variabile E. Desv. var. vidalii (Phil.) LouisMarie, Rhodora 30: 240. 1929. TYPE: Chile. "Ad ostium fluminis Maullin," 41°30'S, Franc. Vidal Gormaz s.n. (holotype, SGO 37066!; isotypes, BAA3424 ex SGO!, BAA-3425 ex SGO!; SGO-PHIL231!, US fragm. & photo ex SGO-PHIL-231!; US photo ex SGO-37066!). Trisetum lechleri (Steud.) Nicora, Fl. Patag. 3: 252. 1978. Basionym: Koeleria lechleri Steud., Syn. PL Glumac. 1: 294. 1854. TYPE: Chile. X Region: Arique, W. Lechler 311 (holotype, P-STEUD-175!; isotype, US fragm. ex P!). Volume 92, Number 4 2005 Finot et al. Trisetum in South America Perennial, sometimes with short rhizomes; culms 30•80 cm tall, glabrous, the upper internode usually very long; nodes 2 or 3, pubescent. Leaf sheaths 60•150 mm long, shorter than the internodes, pubescent; ligule 1•4 mm long, truncate to obtuse, dorsally glabrous or pilose, margins ciliate; blades 5•20 cm X 0.2•5 mm, flat to conduplicate, soft, pubescent. Panicles (4•)6•15 cm X 1^1 mm, contracted to spiciform, interrupted or not, usually dense, the lower branches 2•3 cm long, green, sometimes slightly purple, shining; rachis glabrous to scabrous. Spikelets 5.5•9 mm long, 2- to 4(to 5)flowered, open at the apex; pedicels 1•4 mm long, scabrous; rachilla 0.7•1 mm long, pubescent, the trichomes up to 1.5 mm long; glumes shorter than the florets, unequal, aristulate at the apex or acute; first glume 3•6 X 0.3•0.5 mm, l(or 3)-nerved, linear-lanceolate, attenuate; second glume 4•8 X 0.8• 1.1 mm, 3(or 5)-nerved, oval-lanceolate to oval; florets 6.5•9.5 X 0.6•0.8 mm; lemmas linear-lanceolate, somewhat rounded on the back, glabrous, awned dorsally, dorsally scabrous; apex biaristulate, apical awns up to 1 mm long; awn borne on the upper 1/3 or 1/4 ca. 2•3.5 mm from the apex, geniculate, curved, weakly twisted, scabrous; callus pubescent, the trichomes 0.1•0.5 mm long; paleas 4.5•6.5 mm long, hyaline, 2-nerved, the nerves scabrous; 2-setulate; anthers (0.7•)1.2•1.7 mm long; lodicules 0.6•1.4 mm long, bilobed at the apex, the lobes acute; ovary glabrous. Caryopses ca. 4 X 0.6•0.7 mm, glabrous; endosperm liquid. shorter than the florets and each glume has a short awn at the apex. The first glume is 3.5 mm long, 1-nerved, lanceolate and the second glume is 4 mm long, 3-nerved, oval-lanceolate. The lemma is glabrous, 2-aristulate at the apex, awned on the upper 1/3 and the palea is shorter than the lemma in both florets. On the basis of the characters seen in the type specimens of T. variabile and T vidalii at P, and the original Latin descriptions of the species, it is not possible to distinguish T variabile and T vidalii from T caudulatum. In reference to Deschampsia lasiantha (= Trisetum preslei), T andicola (= Trisetum longiglume), T. andinum, T. biforum (= Trisetum dianthemum), T. hirsutum (= Trisetum phleoides), T. malacophyllum (= Trisetum spicatum var. cumingii), T. chiloense, T ochrostachyum, T monticola, T. variabile, and T. vidalii (all five species = Trisetum caudulatum var. caudulatum), the disposition of the R. A. Philippi collections has been a logistic problem for agrostologists for over 80 years. After her visit to Vienna in 1922, Agnes Chase indicated in an unpublished report (1923, Agnes Chase memorandum to the Bureau of Plant Industry, original at US!) on Edward Hackel's unpublished "Report on Grasses of Chile, 1920" (copy at US!), that Philippi's grass types were loaned to Hackel in Vienna by Philippi via K. F. Reiche (Herb. Mus. Nat. Chile, SGO), after these were returned from US to SCO. The US photographed the entire herbarium and retained fragments of most of the collections including types. Chase (1923, unpublished) noted that Hackel's annotated copy of Reiche's list of the Philippi collections was sent by Hackel to Reiche in Santiago de Chile in 1914. Reiche replied that he was leaving Chile for Mexico, and Hackel never heard from him again. The specimens were never returned to SGO and remain at W stamped "Herb. Hackel" (verified by Bruno Wallnofer, pers. comm. with RJS, 2001). Louis-Marie (1929: 240) created the new name Trisetum variabile var. intonsum to replace the illegitimate T variabile var. virescens (Nees ex Steud.) Macloskie (Macloskie, 1904: 206; not T. variabile var. virescens E. Desv. (Desvaux, 1854: 351)), based on T virescens Nees ex Steud., a plant described for India and placed in Helictotrichon by Henrard (1940) and in Arrhenatherum by Potztal (1968). Thus, T. variabile var. intonsum Louis-Marie is not a species from the New World. Steudel (1853•1855) described Trisetum heteronymum with a variety (Variat: statura et omnibus partibus minoribus, spiculis trifloris, Bromus nr. 117, Bert., Chili). In P, BAA, and SGO there exists a specimen designated as T. heteronymum Steud. Ch romosome num. ber. In = 42 (Baeza et al., 2001). Distribution and habitat. Trisetum caudulatum var. caudulatum grows in Argentina (Neuquen, Rio Negro, and Chubut) and Chile (Region II to XII) in prairies, woods, and heath along roadsides, frequently in sandy soils between 10 and 2200 m. Marticorena et al. (1998) reported the species for Isla Robinson Crusoe (Masatierra), Archipielago de Juan Fernandez, Chile. Phenology. Flowering between October and March. Illustrations. Nicora (1978: 251, fig. 164A-E). Comments. The type of Trisetum caudulatum is represented in the Trinius Herbarium (LE-TRIN) by a fragment containing four spikelets (Nicora, 1978). One spikelet of this type is housed at BAA3364 annotated by Parodi on 31 Jan. 1936, who wrote "La especie esta representada en Leningrado por 4 espiguillas en un pequefio sobre (Nicora, 1978). No trae datos. El nombre bajo el cual figura es T. caudulatum.'''' The spikelet in BAA is 5.5 mm long, contains two florets with glumes unequal, 547 548 Annals of the Missouri Botanical Garden var. minor triflora. This name is not a valid name, de la cumbre, Munoz & Johnson 2583 (SGO); in hills E of Tiltil, Oct. 1958, Bailey s.n. (CONC); Santiago, El Canelo, Gunckel 21997 (CONC); Santiago, Rio Clarillo, Quebrada Las Tinajas, Araya 140 (CONC); Santiago, Quebrada La Plata, Schlegel 4065 (CONC); Santiago, Macuh Gunckel 40532 (CONC); Santiago, Montero 538 (CONC); Santiago, Las Vertientes, Gunckel 44566 (CONC); Farellones, Villagrdn & Mesa 414 (SGO); Santiago, Nido de Aguilas, Penalolen, hacia Casa de Piedra, Munoz & Schick 2558, 2562 (SGO). VI Region: Prov. Colchagua, San Fernando, cerro Echaurren, Montero 1354 (CONC); Agua Buena, al interior de San Fernando, Munoz & Johnson 2635, 2630 (SGO). VII Region: Prov. Linares, La Veguilla, camino de Linares al Melado, Munoz 2709 (SGO); Cajon de Los Cipreses, von Dessauer s.n. (SGO); Prov. Linares, Valle Gualquivilo, Banos de Azufre, Schlegel 3645 (CONC). VIII Region: Prov. Concepcion, La Toma, 29 Oct. 1934, Junge s.n. (CONC); Concepcion, Nov. 1896, Neger s.n. (fragm. CONC-148139 ex M); Concepcion, 10 Oct. 1951, Pfister s.n. (CONC); Concepcion, Barros 1987 (CONC); Talcahuano, Parque Hualpen, Carrasco 266 (CONC); San Pedro, Oct. 1943, Pfister s.n. (CONC); San Pedro, Ricardi, Marticorena & Torres s.n. (CONC); San Pedro, 12 Nov. 1944, Pfister s.n. (CONC); San Pedro, 10 Dec. 1946, Pfister s.n. (CONC); Concepcion, a orillas de camino Concepcion•Florida, Matthei 174 (CONC); Concepcion, Cerro Caracol, Pfister 370 (CONC); Bio-Bio, Hacienda Las Canteras, entre Antuco y Los Angeles, Munoz & Johnson 2737 (SGO); Entre Antuco y Tumbunleo, Munoz & Johnson 2732 (SGO); La Laja, Malacura, Munoz & Schick 1493 (SGO); Parque Nacional Laguna del Laja, Antuco, Finot & Baeza 1 (CONC); Camino Canteras•Antuco, 20 km antes de Antuco, Finot & Baeza 2 (CONC); Malalcura, cerros, Finot & Baeza 7 (CONC); Parque Nacional Laguna del Laja, Los Barros, sector Aduana, Finot & Baeza 14 (CONC); entre Chacay y canchas de Sky, Finot & Baeza 2074 (CONC); cercanias de Rere, Yumbel, Munoz 2717 (SGO); Isla Quiriquina, Gunckel 13797 (CONC); Nuble, camino a las Termas, 30 km de Chilian. Munoz & Johnson 2676 (SGO); Atacalco, 26 Nov. 1944, Pfister s.n. (CONC); 5 km E de Quilldn, Hutchinson 212 (SGO, F); Concepcion, 1 Nov. 1927, Barros s.n. (CONC); Tome, 24 Nov. 1925, Barros s.n. (CONC); Lirquen, 11 Dec. 1950, Ricardi s.n. (CONC); Tome, 14 Nov. 1925, Barros s.n. (CONC). IX Region: Malleco, Lumaco, Santa Clara, Gunckel 630 (CONC); Laguna Galletue, Montero 4956 (CONC); Quechumalal, Pampa del Olvido, Mar. 1958, Schlegel s.n. (CONC); Valle de Lonquimay, Pfister s.n. (CONC); Cautin, Temuco, Cerro Nielol, Montero 4508 (CONC); Malleco, Parque Nacional de Nahuelbuta, Piedra del Aguila, Ricardi, Marticorena & Matthei 1976 (CONC); Termas de Tolhuaca, Gunckel 16027 (CONC); Prov. Cautin, Temuco, Padre Las Casas, Montero 1962 (CONC); Cautin, Volcan Llaima, Gunckel 15101 (CONC); Villarica, Tromen, Limite Chileno-Argentino, Ricardi & Matthei 27 (CONC). X Region: Valdivia, Jan. 1860, R. A. Philippi s.n. (SGO); Llanquihue, Puerto Varas, Petrohue, Marticorena, Weldt & Crisci 1982 (CONC); Rio Palena, Jan. 1887, Delfin s.n. (SGO); Chiloe, Lago Rio Negro, Villagrdn, Aguila & Leiva 6954 (CONC); Valdivia, San Juan, Jan. 1855, R. A. Philippi s.n. (SGO); Lago Llanquihue, without date, Philippi s.n. (SGO); Puerto de Corral, Munoz & Johnson 3211 (SGO); Puerto Montt, Feb. 1858, Philippi s.n. (SGO); Puerto Montt, Fonck 71 (SGO). XI Region: Aisen, Cohaique, Schlegel 2371 (CONC). XII Region: Tierra del Fuego, San Sebastian, Ricardi & Matthei 236 (CONC); Caleta Josefina, Ricardi & Matthei 166 (CONC). and the specimen is Bertero 116, not Bertero 117 as the protologue indicates. These specimens were determined to be synonyms of T. caudulatum. Additional specimens studied. ARGENTINA. Chubut: Lago Futalaufquen, en la cascada, Krapovickas 3933 (SI); P.N. Los Alerces, Lago Cisne, Roquero 452 (BA); Carrenleufu, lllin s.n. (LP); Pampa Chica, Soriano 2490 (BAA); Lago Futalaufquen, claros del bosque de coigiie y cipres, Burkart 19905 (BAA); PN. Los Alerces, Rio 2, Lahitte & Roquero 363 (BA); Corcovado, lllin s.n. (LP); Cerro Leleg, 14 Jan. 1949, Perez-Moreau s.n. (BA). Neuquen: Lago Guillen, 28 Dec. 1937, Kalela s.n. (S); San Martin de Los Andes, Bridarolli 2147 (LP 39146); San Martin de Los Andes, Dawson 1285 (LP); PN. Lanin, Lago Lacar, camino a Angostura, 17 Feb. 1961, Leon & Calderon s.n. (BAA); Isla Victoria, Corte 34 (LP); Nahuel Huapi, Puesto Panuelo, 18 Jan. 1930, Offermann s.n. (BAA); Fuerte Chacabuco, Parodi 15611 (BAA); Parque Nacional Lanin, Arroyo Grande, S.L. Lacar, Eskuche & Klein 316 (CTES); Parque Nac. Lanin, Pampa Hui Hui, Eskuche & Klein 1417-27 (CTES); Nahuel Huapi, F. Chacabuco, Vellerini 297 (BAA); Los Lagos, Villa La Angostura, Lago Nahuel Huapi, bahia proxima a Peninsula Cumelen, Rugolo 1188 (CONC); Los Lagos, Villa La Angostura, cerro Bayo, Rugolo 1283 (CONC); Lago Epulafquen, Dawson & Schwabe 2481 (BAA); P.N. Lanin, Lago Paimiin, Lahitte, Roquero & Lopez 486 (BA); P.N. Lanin, San Martin de Los Andes, Lahitte, Roquero & Lopez 67 (BA); P.N. Lanin, San Martin de Los Andes, Lago Lacar, Quila-Quina, Roquero 326 (BA); P.N. Lanin, Lago Lacar, Pucara camino a Angostura, Leon & Calderon 1285 (BAA). Rio Negro: Lago Gutierrez, 27 Nov. 1937, Kalela s.n. (S), Lago Gutierrez, Kalela 1277 (S); Puerto Moreno (Los Juncos), San Ramon, Loma Grande, Vellerini 265 (BAA); Puerto Moreno (Los Juncos), Ea. San Ramon, Vellerini 257 (BAA); 6 km from Bariloche, Pedersen s.n. (CTES); Leleque, Soriano 2409 (BAA); Catedral Hotel, P.N. Nahuel Huapi, Pedersen 1468 (CTES); P.N. Nahuel Huapi, Boelcke & Correa 5864 (BAA); picada a lo largo de Martin Grande, Boelcke & Correa 6142 (BAA); Bariloche, arroyo Guillelmo, camino a El Bolsdn, Nicora 7478 (BAA). CHILE. II Region: El Rincon, just N of Paposo, along trail to old Paranas Mine, Johnston 5539 (BAA); Antofagasta, Dpto. Taltal, vie. of Aguada de Miguel Diaz, Johnston 5407 (SGO). IV Region: Ovalle, bosque de Talinay, lado sur de la desembocadura del Limari, Munoz & Coronel 1232 (BAA); Limari, Cordillera de Ovalle, Tulahuen-Leiva, Jiles 4694 (CONC). V Region: Valparaiso, In sylvaticus calidis collium loco dicto La Laguna, Valparaiso, 1829, Bertero 997 (P, BAA ex P, SGO ex P); cerro Roble above Calco, Hutchinson 59 (SGO); Valparaiso, Lechler 2846 (US fragm. ex P, F); Cuesta La Dormida, camino entre Quillota y Santiago, Munoz & Schick 1545 (SGO); Aconcagua, Petorca, carretera Panamericana, 5 km N de Longotoma, Ricardi, Marticorena & Matthei 1829 (CONC); Quillota, Limache, Cerro Cruz, Garaventa 2248 (CONC); Quillota, cerro La Campana, Zoellner 13135, 18030 (CONC); Cocalan, 9 Nov. 1913, Baeza s.n. (CONC); Archipielago de Juan Fernandez, Oct. 1872, F. Philippi s.n. (SGO); Quillota, Limache, cerca del pueblo, Garaventa 6457 (CONC); Valparaiso, Cerro La Campana, Hutchinson 45 (US); Cerro Las Vizcachas, Hutchinson 110 (SGO); Quebrada Verde, Munoz & Johnson 2524 (SGO). Region Metropolitans: Cuesta de Chacabuco, a 3 km Volume 92, Number 4 2005 8b. Trisetum caudulatum Trin. cora, Fl. Patag. 8(3): 254, TYPE: Argentina. Chubut: Futalaufquen, A. Soriano BAA-3365!). Finot et al. Trisetum in South America var. correae Nifig. 164e. 1978. Futaleufu, Lago 4334 (holotype, Ovary and caryopses with trichomes at the apex. Distribution and habitat. Nicora (1978) described this variety on the basis of material collected in Neuquen, Rio Negro, and Chubut, Argentina. In Chile this plant grows between Region V and Region X generally between 1000 and 2000 m. Rodulfo Amando Philippi appears to be the first to have collected this species in Region X, Chile (Finot, 2002). Trisetum caudulatum var. correae grows along the Andes and is a forage species. Phenology. Flowering between December and March. Additional specimens studied. ARGENTINA. Chubut: P.N. Los Alerces, Lago Futalafquen, entre Rio Desaguadero y Arroyo del Salto, Lahitte & Roquero 277 (BA); Lago Futalaufquen, 7 Ian. 1964, Lahitte s.n. (BA). Neuquen: Dpto. Los Lagos, 3 km W de Confluencia, Gonzalez 766 (LP); Lago Traful Sur, orilla del lago, Riigolo de Agrasar 233 (BAA); P.N. Nahuel Huapi, Lago Trafuel, El Mirador, Boelcke & Hunziker 3657 (BAA); P.N. Nahuel Huapi, Campamento Rio Villegas, 29 Jan. 1941, PerezMoreau s.n. (BAA, paratype). Rio Negro: P.N. Nuahuel Huapi, faldeos cerro Santa Elena, Fabris & Solbrig 1171 (LP). CHILE. V Region: Punta Iman, Cerro Roble, Zoellner 18196 (ZOELLNER). IX Region: Valle de Lonquimay, 5 Jan. 1947, Pfister s.n. (CONC). X Region: Lago Llanquihue, without date, Philippi s.n. (SGO); Posada del Valle, Jan. 1877, Philippi s.n. (SGO). 9. Trisetum dianthemum (Louis-Marie) Finot, Contr. U.S. Natl. Herb. 48: 664. 2003. Basionym: Trisetum spicatum var. dianthemum Louis-Marie, Rhodora 30: 239. 1929. TYPE: Chile. X Region: Provincia de Llanquihue, grama a orillas del Rio Puelo, 1872, F. Vidal Gormaz s.n. (holotype, SGO-PHIL-239b!; isotypes, BAA-3358!, SGO-37069!, SGO-68170 photo!, US fragm. ex SGO-PHIL-239b & photo!, US photo ex SGO-37069!). Perennial, caespitose; culms 18•60 cm tall, geniculate, sericeous or pilose below the panicle; nodes 1 or 2, nearly basal, glabrous. Leaf sheaths 3^1 cm long, glabrous; ligule 1•1.5 mm long, membranous, truncate to obtuse, ciliate-denticulate, dorsally glabrous; blades 5•15 cm X 1.5•2.5 mm, glabrous, smooth, scabrous or ciliate on the margins. Panicles 4•8 X 1•2 cm, somewhat lax, subspiciform, shining; rachis pubescent. Spikelets 6.5• 8 mm long, 2- or 3-flowered, open at the apex; pedicels 1.5^1 mm long, scabrous; rachilla 0.9•1 mm long, pubescent, the trichomes 0.2•1 mm long; 549 glumes longer than the florets, subequal, lanceolate, the first glume nearly as long as and slightly narrower than the second glume; keel scabrous; margins hyaline; apex aristulate; first glume 6•7.5 X 0.6•0.8 mm, l(3)-nerved; second glume 6.5•8 X 0.8-1 mm, 3-nerved; florets (first) 5.6-6.3 X 0.5•0.7 mm, upper floret ca. 5.5 mm long; lemmas glabrous, slightly scabrous on the keel, dorsally awned; apex hyaline, 2-setulate; margins hyaline; awn 3.5•7 mm long, borne on the upper 1/3 or 1/4, divaricate, weakly twisted, scabrous; callus obtuse, with a few trichomes ca. 0.2 mm long; paleas 4.2• 4.5 mm long, shorter than the lemma; apex 2-setulate, 2-nerved, the nerves scabrous; anthers ca. 0.8 mm long; lodicules ca. 0.6•0.8 mm long; apex 3-lobed; ovary glabrous. Caryopses not seen. Distribution and habitat. Endemic to southern Chile and Argentina, where it is known only from Region X in Chile and from Chubut, Argentina, between 3 and 210 m. Comments. Louis-Marie (1929) established the new name Trisetum dianthemum for the illegitimate T. biflorum Phil. (Philippi, 1873: 568) and simultaneously placed it in varietal rank (T. spicatum var. dianthemum (Phil.) Louis-Marie). Trisetum dianthemum differs from T spicatum (L.) K. Richt. by having glumes longer than the florets (vs. both glumes shorter than the florets in T spicatum), longer spikelets (6.5•8 mm long in T dianthemum vs. 4.5•6 mm in T. spicatum), isomorphic glumes (vs. subequal glumes in T. spicatum), and panicles somewhat lax, and somewhat spiciform (vs. spiciform in T. spicatum). Phenology. Flowering between December and January. Additional specimens studied. ARGENTINA. Chubut: Lago Los Nifios, Nicora 9610 (SI). CHILE. X Region: Puerto Varas, Islote frente a Punta Guano, Marticorerw, Weldt & Crisci 1967 (CONC); Prov. Valdivia, Barra del Rio Bueno, Hollermayer 1252 (CONC, LP); Quefii, Jan. 1887, 0. Philippi s.n. (BAA-3357 fragm. ex SGO, SGO, US photo ex SGO); Panguipulli, 14 Dec. 1927, Hollermayer s.n. (CONC); Panguipulli, Hollermayer 26-a (CONC). 10a. Trisetum longiglume Hack. var. longiglume, Repert. Spec. Nov. Regni Veg. 7: 319. 1909. TYPE: Argentina. Mendoza: "in monte Piedra del Burrero prope San Rafael," Jan. 1897, E. Wilczek 571 (holotype, W not seen). Trisetum andicola Louis-Marie, Rhodora 30: 244. 1929. Deschampsia andicola (Louis-Marie) Valencia, Revista Argent. Agron. 8: 129, f. 2. 1941. TYPE: Chile. Santiago: Laguna Negra, 2700-4000 m, Mar. 1873, F Vidal Gormaz 265 (holotype, US-556459 fragm.& 550 Annals of the Missouri Botanical Garden photo ex SGO-PHIL-265!; isotypes, BAA-3344 fragm.!, SGO-37123!, SGO-37124!). Perennial, caespitose; culms 20•30 cm tall, glabrous, erect or geniculate at the base; node 1, basal. Leaf sheaths 2•6 cm long, glabrous; apex with sheath auricles; ligule 0.2•2 mm, dorsally glabrous, denticulate at the apex; blades 3.5•12 cm X 1.5• 2 mm, flat, glabrous, abaxially smooth, slightly scabrous on adaxial surface and margins. Panicles 3.5•6 X 1•3 cm, subspiciform, ovoid, dense, green to pale purple, the branches ascending; rachis glabrous; pedicels ca. 3 mm long, glabrous. Spikelets ca. 9 mm long, 2- or 3-flowered; rachilla 1.5 mm long, conspicuously pubescent, the trichomes up to 4 mm long; glumes usually longer than the florets, lanceolate, isomorphic, slightly tinged with green and purple; keel mostly smooth except near apex; apex attenuate, somewhat aristulate; first glume 8.5•9.5 X 0.6•0.7 mm, 1- to 3-nerved; second glume 8.5•9.5 X 0.8•1.2 mm, 3-nerved; florets 5• 6.2 mm long; lemmas lanceolate, laterally compressed, dorsally awned, glabrous, slightly keeled, 5-nerved; apex 4-aristulate, apical awns are projection of the intermediate nerves; marginal apical awns hyaline, shorter than the middle apical awns: awn 8•9.5 mm long, borne near the median, 2.5• 3 mm from the apex, twisted at base; callus obtuse, with trichomes 2•3.5 mm long, about 2/3•3/4 as long as the lemma; paleas 5•5.3 mm long, shorter than the lemma, hyaline, 2-nerved, the nerves scabrous; apex bidentate; anthers (0.5•)0.8•1 mm long; lodicules 0.6•0.8 mm long, hyaline, tridentate at the apex, the teeth small, the middle tooth a little larger than the lateral teeth; ovary pubescent, with a few trichomes near apex. Caryopses 2.0•2.7 mm long, rostrate, pubescent, apex with a few short curved and shining trichomes; trichomes curved and shining; hilum short, ovate; endosperm soft. Distribution and habitat. Trisetum longiglume var. longiglume is endemic and rare to south-central Chile (Metropolitana) and Argentina (Mendoza, Neuquen, and Chubut) between 2000 and 4000 m. This species is primarily found in Andean valleys. Phenology. Flowering between January and March. Comments. Louis-Marie (1929) described Trisetum andicola on the basis of material collected in the Cordillera de Los Andes of the Region Metropolitana in Chile. Valencia (1941) transferred the taxon to Deschampsia. Later Parodi (1949b) thought it should be treated again as Trisetum. This species shows unique characters, making it easily recognized: glumes isomorphic, longer than the spikelet, the lemmatal awn borne near the middle of the lem- ma, long trichomes on the callus, and, in the typical variety, trichomes at the apex of the ovary and on the caryopsis. Additional specimens studied. ARGENTINA. Chubut: Mt. La Torta, Rivadavia Range, 30, Beetle & Soriano HS-381a (US). Neuquen: Dpto. Minas, Paso del Macho, Boelcke et al. 13932 (BAA); Copahue, 3 Jan. 1930, Hutchinson s.n. (BAA). CHILE. Region Metropolitana: Cajon Las Lefias, Arroyo et al. 94447, 94454 (CONC). 10b. Trisetum longiglume var. glabratum Nicora, Fl. Patagonica 3: 245, f. 158. 1978. TYPE: Argentina. Neuquen: Dpto. Lacar, Estancia Meliquina, Co. Repollo, Z. E. Rugolo de Agrasar & E. Agrasar 573 (holotype, BAA!). Perennial, with short rhizomes; culms JO•38 cm tall, erect, glabrous. Leaf sheaths glabrous; ligule ca. 2 mm long, glabrous, minutely denticulate at the apex; blades stiff, flat to conduplicate, abaxially glabrous, adaxially with JO to J2 prominent, scabrous ribs; margins scabrous; lower blades 5•6 cm X J•2 mm; upper (flag) blade J cm long. Panicles few-flowered (J2 to 2J spikelets), linear, narrow; rachis glabrous; pedicels scabrous. Spikelets ca JO mm long, 2-flowered; rachilla 2•2.5 mm long, pilose, the trichomes up to 3 mm long; glumes ca. JO mm long, as long as the spikelet, isomorphic; first glume J- to 3-nerved; second glume 3-nerved; florets 6.5•8.6 mm long; lemmas dorsally awned, glabrous, 5-nerved, the median and marginal nerves very conspicuous toward the apex, the nerves projected as 4 short apical awns; apical awns J•2 mm long; awns 8•JJ mm long, borne near the median, geniculate and twisted; callus very pubescent, the trichomes J/2 the length of the lemma; paleas hyaline, a little shorter than the lemma; anthers J.3• J.7 mm long; ovary glabrous. Caryopses 3.5•4.2 mm long; hilum oval. Distribution and habitat. Trisetum longiglume var. glabratum is endemic to Argentina (Neuquen and Chubut) (Nicora, J978; Zuloaga et al., J994). This species occurs in rocky soils in the high Andes. Illustrations. Nicora (J978: 247: fig. J58A-E). 11. Trisetum macbridei Hitchc, Contr. U.S. Natl. Herb. 24(8): 359. J927. TYPE: Peru. Huaron, collected on rocky NE slope, 4200 m, J2 June J922, /. F. Macbride & Featherstone 1131 (holotype, US-JJ6J5J0!; isotypes, F050J68!, F-5J7642!, P!, S-fragm.!). Perennial, with short rhizomes; culms 20^10 cm tall, glabrous, erect; nodes J or 2, basal. Leaf Volume 92, Number 4 2005 sheaths pilose; apex with sheath auricles as long as ligule; ligule 1•1.5 mm long, truncate-obtuse, denticulate, ciliate, dorsally pilose; blades 2•10 cm X ca. 3 mm, long, flat to conduplicate, stiff, sparsely pilose abaxially, scabrous adaxially, ciliate on the margin; upper blade 2•3 cm long. Panicles 6•8 X ca. 1 cm, spiciform, narrow, green-purple, shining; rachis glabrous. Spikelets ca. 8 mm long, 2-flowered; pedicels distally scabrous; rachilla ca. 1.5 mm long, pubescent, the trichomes ca. 1 mm long; glumes exceeding the florets by 1/3•1/2 in length, equal to subequal, ovate; keel somewhat scabrous; apex acute, aristulate, scabrous; first glume 7.5•8 X 1.1•1.5 mm, 1-nerved; second glume 7.8•8 X 1.3•1.5 mm, 3-nerved; florets ca. 4•5.5 X 0.9 mm; lemmas glabrous, green on back, somewhat purple on margins, dorsally awned; apex 2-aristulate, the apical awns 0.4•0.5 mm long; awn borne on the upper 1/3, twisted and geniculate, a little scabrous; callus obtuse, with trichomes 0.1•1 mm long; paleas 3 mm long, 1/2•2/3 as long as the lemma, hyaline, 2-nerved, the nerves scabrous; apex erose, minutely ciliate; anthers ca. 1 mm long, ovate; lodicules 0.6•0.7 mm long, with 2 or 3 small lobes at the apex, one of the lobes larger than the other two, sometimes cleistogamous flowers with anthers ca. 0.5 mm long; ovary glabrous. Caryopsis not seen. Distribution and habitat. Trisetum macbridei is an endemic and rare species found only in the Andes of Central Peru. Tovar (1993) reported T. macbridei from Huancavelica and Pasco, between 4200 and 4500 m. This species is found on rocky slopes. Phenology. Flowering in June. Comments. Trisetum macbridei is related to T spicatum (Hitchcock, 1927), from which it differs in having both glumes conspicuously longer than the florets (vs. glumes shorter than the florets in T spicatum), glumes similar in shape (vs. glumes dissimilar in T. spicatum), culm glabrous below the panicle (vs. culm pubescent or pilose below the panicle in T spicatum), and leaves pubescent (vs. glabrous in typical T spicatum). On the basis of the isomorphic glumes, equal in length and width, Louis-Marie (1928, 1929) classified this species in Trisetum subgenus Isolytrum. Additional specimens studied. PERU. Ancash: Huaylas, Huascaran National Park, quebrada Alpamayo above Lago Jancarurish, Smith, Valencia & Gonzdles 9772 (F). Jumn: Prov. Cerro Huaron, rocky lakeshore, Asplund 11784 (S), 11793 (S, US). 12. Trisetum mattheii Finot, sp. nov. TYPE: Chile. Region I: Tarapaca, camino de Arica al Portezuelo de Chapiquifia, km 111, 18°18'S, 69°30'W, 4100 m, 9 Feb. 1964, C. Martico- Finot et al. Trisetum in South America 551 rena, 0. Matthei & M. Quezada 86 (holotype, CONC-88160!). Figure 1. Gramen caespitosum; culmi 20•36 cm alti, erecti, pilosi; vaginae inferiores pilosae, superiores glabrae; ligula hyalina, triangularis, dentato-ciliata, 1.5•2 mm longa; laminae conduplicatae, inferiores pilosae, superiores glabrae; panicula spiciformis 3•5.5 X 0.5•0.8 cm, pauciflora; spicula 2-flora, 4.5•5 mm longa; glumae inaequales, inferior angustior, 1-nervia, 3.5•4.5 X 0.5•0.6 mm, lineari-lanceolata, superior 3-nervia, ovato-lanceolata, 3.8• 4.7 X 0.7•0.9 mm; lemma hirsuta, aristata ad 1/3 superiorem; arista divaricata, non torta, scabra; callus obtusus, pilosus, pilis 0.2 mm longis; palea brevior quam lemma, hialina, binervata, nervis scabris, bidentato-biaristulata ad apicem; lodiculae 0.6•0.8 mm longae, bilobulatae ad apicem, lobulis acutis; stamina 3, antheris 1.1 mm longis; ovarium apice glabro. Perennial, caespitose; culms 20•36 cm tall, purple, pubescent below the panicle, the trichomes ca. 0.5 mm long, first antrorse, then retrorse; nodes 2, glabrous. Leaf sheaths 1•4 mm long, longer than the internodes; lower sheaths pilose, with age glabrous; culm sheaths glabrous; ligule 1.5•2 mm long, subtriangular, dentate and minutely ciliate at the apex; dorsally glabrous; blades 1•6 cm X 0.5• 1 mm, conduplicate, almost filiform, somewhat stiff, scabrous on the margin and adaxial surface; lower blades pilose; upper blades glabrous. Panicles 3• 5.5 X 0.5•0.8 cm, tinged with purple and yellow, spiciform, narrow, linear, few-flowered, sometimes interrupted at the base, acute at the apex; rachis densely pubescent, the trichomes ca. 0.5 mm long. Spikelets 4.5•5 mm long, 2-flowered; pedicels densely pubescent; rachilla 0.6•0.9 mm long, with stiff trichomes at the base, glabrous toward the apex; beyond upper floret rachilla with an aristiform appendix up to 0.5 mm long; glumes a little shorter than the spikelet, dissimilar; apex acute, sometimes short-awned; first glume 3.5•4.5 X 0.5• 0.6 mm, linear-lanceolate, 1-nerved; second glume 3.8•4.7 X 0.7•0.9 mm, ovate-lanceolate, 3-nerved; florets 3.5^1 X 0.6•0.7 mm; lemmas green at the base, purplish toward the apex, awned dorsally, hirsute, the trichomes 0.3•0.5 mm long; apex 2-aristulate, the apical awns ca. 0.3 mm long; awn 2.5• 3.5 mm long, borne on upper 1/3, 1•1.5 mm from the apex, divaricate, not twisted nor geniculate, scabrous; callus pubescent, with trichomes ca. 0.2 mm long; paleas 2.8•3 mm long, shorter than the lemma, 2-nerved; keels scabrous below; apex 2dentate, the teeth prolonged as hyaline setae; anthers ca. 1.1 mm long; lodicules 0.6•0.8 mm long, deeply bilobed at the apex; ovary glabrous. Caryopses not seen. Distribution and habitat. K nown on.ly fn the 552 Annals of the Missouri Botanical Garden Figure 1. Trisetum mattheii. •A. Habit. •B. Sheath, ligule, and portion of the blade. •C. Spikelet. •D. Floret. •E. Rachilla and palea, dorsal view. •F Lodicules and palea, ventral view. •G. Rachilla. •H. Lodicules. •I. Caryopsis. Marticorena, Matthei & Quezada 86 (holotype, CONC). Volume 92, Number 4 2005 Finot et al. Trisetum in South America type locality, in Cordillera de Los Andes, northern Chile, Region I, at 4100 m. Phenology. Flowering in February. Comments. Trisetum mattheii is allied to T. oreophilum Louis-Marie var. johnstonii Louis-Marie, and can be separated from the latter by having taller culms (20•36 cm), with linear, narrow and fewflowered panicles, glabrous sheaths and upper blades, and subtriangular ligules. Trisetum oreophilum var. johnstonii is smaller (10•20 cm tall), with densely flowered, ovate panicles, pubescent sheaths and blades, and truncate ligules. Etymology. The specific epithet honors the Chilean agrostologist, Oscar Matthei Jensen, retired Professor at the Universidad de Concepcion, Chile. to shortly awned, the awn up to 1 mm long; second glume 5.3•7.6 X 0.7•1.1 mm, oval-lanceolate, awned at the apex, 3-nerved; keel scabrous; florets 6.5•7.5 X 0.6•0.7 mm; lemmas dorsally awned, hirsute, green dorsally; margins and apex hyaline, tinged with purple; apex biaristulate, the awns ca. 1 mm long; awn 6•9 mm long, borne on the upper 1/3, 1.5•2.5 mm below the apex, geniculate, not twisted; paleas 4•5 mm long, shorter than the lemma, hyaline; apex biaristulate; anthers ca. 1.5 mm long, yellow; lodicules 0.8•1 mm long, apex bilobed, the lobes acute; ovary glabrous. Caryopses 2.5•3 X 0.5•0.6 mm, glabrous; endosperm liquid. 13. Trisetum nancaguense Finot, sp. nov. TYPE: Chile. Region VI: Prov. Cardenal Caro, 12 km E of Pichilemu on Hwy. toward Nancagua, 34°23'S, 71°59'W, 45 m, 15 Nov. 1990, T G. hammers, C. M. Baeza y P. Penailillo 7894 (holotype, CONC-113221!; isotype F2183048!). Figure 2. Gramen caespitosum, 40•75 cm altum; culmus erectus vel geniculatus, glaber, folia multo excedens; vaginae laxae, pilosae; ligula 2•3 mm longa, hialina, truncata, dentato-ciliata; laminae 4•15 cm X 2•5 mm, planae, pilosae; panicula 6.5•9 X 0.8•2 cm, contracta; spicula 5.5•8 mm longa, 2- v. 3-flora, lateraliter compressa; rhachilla 1 mm longa, dense pilosa; glumae inaequales; inferior gluma 3.5•6 X 0.2•0.4 mm, lineari-lanceolata, 1-nervata, angustior; superior gluma 5.3•7.6 X 0.7•1.1 mm, ovato-lanceolata, 3-nervata, apice acuminata; lemma hirsuta, aristata ad 1/3 superiorem; arista flexuosa vel geniculata, non torta, scabra; callus obtusus, pilosus; palea brevior quam lemma, hialina, binervata, nervis scabris, bidentato-biaristulata ad apicem; stamina 3, antheris 1.5 mm longis; lodiculae 0.8•1 mm longae, bilobulatae ad apicem, lobulis acutis; ovarium apice glabro; caryopsis 2.5•3 X 0.5•0.6 mm, glabra; endospermium liquidum. Perennial, caespitose; culms 40•75 cm tall, glabrous; nodes 2 to 4, glabrous or pilose. Leaf sheaths shorter than the internodes, pubescent; lower sheaths 3•8 cm long; upper sheaths 15•20 cm long; ligule 2•3 mm long, truncate, dentate-ciliate; blades 4•15 cm X 2•5 mm, flat, soft, pubescent abaxially, sparsely pilose adaxially. Panicles 6.5•9 X 0.8•2 cm, subspiciform, contracted, sometimes interrupted at the base, silvery-green to weakly purple; rachis scabrous. Spikelets 5.5•8 mm long, 2- or 3-flowered; pedicels 2•3.5 mm long, pilose to scabrous; rachilla ca. 1 mm long, pilose, the trichomes 2•3 mm long; glumes dissimilar, shorter than the florets, greenish; margins membranous, margins and apex tinged with purple; first glume 3.5•6 X 0.2•0.4 mm, linear-lanceolate, subulate, 1-nerved; keel scabrous on upper half; apex acute 553 Distribution and habitat. Trisetum nancaguense is endemic to Chile, ranging from Region Metropolitana to VIII Region, between 33°20'S and 36°50'S and from 45 to 2450 m altitude. Phenology. Flowering in November and February. Comments. Trisetum nancaguense appears related to T. barbinode Trin., from which it differs in having pubescent blades (vs. glabrous), leaf sheaths shorter than the internodes (vs. longer), glumes shorter than the florets (vs. equaling or longer than the spikelet), first glume 1-nerved (vs. 1- or 3nerved), lemma hirsute, i.e., covered with short and stiff trichomes (vs. lemmas pubescent, i.e., covered by short and softer trichomes), and by its distribution in the Central Valley (Depresion Intermedia) and coast in Central Chile at low elevations (T barbinode occurs in the Cordillera de Los Andes, usually above 1000 m). Paratypes. CHILE. Region Metropolitana: Cordillera, Valle Nevado, E. Bayer 4608 (CONC). VII Region: Maule, Cerro al SW de Coronel de Maule, G. L. Stebbins 9061 (SGO). VIII Region: Concepcion, camino entre Concepcion y Bulnes km 42, Villarroel & Weldt 151 (CONC). 14a. Trisetum oreophilum Louis-Marie var. oreophilum, Rhodora 30: 221. 1929. TYPE: Peru. Cuzco: moist grassland, high up ravine above Olloutaytambo, 3600 m, 5 Dec. 1923, A. S. Hitchcock 22535 (holotype, US1164163!). Perennial, caespitose; culms 15•60 cm tall, puberulent to densely pubescent below the inflorescence; nodes 2, glabrous. Leaf sheaths shorter than the internodes; lower sheaths pubescent; upper sheaths glabrous; ligule 2^1.5 mm long, dorsally glabrous, denticulate at the apex; blades flat, glabrous, usually with long trichomes near the base; lower blades 8•16 cm X 2 mm; upper leaf blades 3.5-5 cm long. Panicle (5-)7-ll X 0.8-1.5 cm, 554 Annals of the Missouri Botanical Garden Figure 2. Trisetum naneaguense. •A. Habit. •B. Sheath, ligule, and portion of the blade. •C. Spikelet. •D. Floret. •E. Palea, dorsal view. •F. Palea, ventral view. •G. Rachilla. •H. Lodicules. •I. Caryopsis. hammers. Baeza & Penailillo 7894 (holotype, CONC). Volume 92, Number 4 2005 Finot et al. Trisetum in South America subspiciform to spiciform, linear, narrow, denselyflowered, interrupted at the base, purplish-green, shining, rachis pubescent. Spikelets 3.5•4.5 mm long, 2- or 3-flowered, purple; pedicels pubescent; rachilla ca. 0.7 mm long, with long trichomes at the base; glumes unequal, shorter than the florets, dorsally green and purple; margins hyaline; apex acute; first glume 2.2•3.6 X 0.4•0.6 mm, narrower and usually shorter than the second glume, 1nerved; second glume 2.8^1.1 X 0.6•0.9 mm; florets 3.5•4.6 mm long; lemmas pubescent to sparsely pubescent, dorsally awned; margins hyaline; apex 2-aristulate; callus obtuse, with short trichomes; awn 1.5^1 mm long, scabrous, borne on the upper 1/3, 1•1.5 mm below the apex, not twisted nor geniculate, curved, scabrous, sometimes with short trichomes at the base; paleas 2.8•3 mm long, shorter than the lemma, hyaline, 2-nerved, the nerves scabrous; apex 2-dentate; anthers 1•1.2 mm long, yellow; lodicules ca. 0.6 mm long; apex bilobed, hyaline; ovary glabrous. Caryopses ca. 2.5 mm long, glabrous; endosperm liquid. Judziewicz, King & j0rgensen 9176 (US); 9 km NE of San luan de Vefasco on rd. to Lago Colta, Peterson, Judziewicz, King & j0rgensen 9239 (US); Urbina, tow. Mt. Chimborazo, Asplund 7896 (S). Cotopaxi: Lag. Quilotoa, Laegaard 101347 (QCA); EN. Cotopaxi, Limpiopungo, Roig J. 12000 (MERL). Pichincha: Sodiro 1893 (US-1163177, paratype); N slope of Mount Corazon, Asplund 17550 (S); San luan, tow. Chiriboga, Asplund 16131 (S); slope of the Pichincha above Lloa, Asplund 7547 (F, S); Quito, Panecillo, Asplund 6023 (F). PERU. Ancash: Huari, Huascaran Natl. Park, Quebrada Rima Rima, a lateral valley of Quebrada Carhuazcancha, Smith, Valencia, Gonzalez & Buddensiek 12224 (F); Huaylas, Huascaran Natl. Park, Quebrada Los Cedros, Smith, Valencia & Minaya 9943 (F). Cajamarca: Celendin, lalca de Kumulca, Sagdstegui, Mostacero & Leiva 12056 (F); Lagunas Maqui-Maqui, Sdnehez-Vega & Cabanillas 6790 (F); Hualgayoc, a 31 km de Bambamarca, arriba de la ciudad de Hualgayoc, sobre la ladera sur, Sanchez-Vega, Molau & Ohman 3800 (F); Celendin, Quebrada de Sendamal, sobre la carretera a Celendin, Sanchez-Vega, Molau & Ohman 3812 (F); Sexcemayo, cerro Mahoma al W de Cajamarca, Sanchez-Vega & Castillo 6370 (F); Cajamarca-Choten, en el arboretum de CICAFOR, a 4 km de la carretera Pacasmayo•Cajamarca, desvio la altura del km 155, Sanchez-Vega, Torrel & Medina 2557 (F); Cajamarca, a la altura del Paso El Gavilan, Sanchez-Vega et al. 1380 (F). Cuzco: north of city, Hitchcock 22471 (US); Rodadero, Marin 1432 (US); cerro, Vargas 7045 (US); Urubamba, large eroded rock called Maranqaqa on Inca plaza called Capellanpampa, Davis et al. 1381 (F). Huancavelica: Prov. Huancavelica, entre Huaytanayocc y Manta, Tovar 2539 (US); Prov. Tayacaja, Hacienda Huari, Salaverry 17 (US); Hacienda Alalay, entre Mejorada y Pampas, Tovar 2474 (US); Prov. Castrovirreina, Choclococha, Tovar 2933 (US). Junin: Rio Blanco. Swollen 7064 (US); Res. Nac. Jumn, Ondores, Pettersson 41, 57 (S); Goyllarisquisca, NW part of Jumn, Hitchcock 22323 (US-1164158, paratype); Cerro de Pasco, Hitchcock 22254 (US-1164159, BAA-3408, paratype); La Quinhua, Cerro de Pasco, Hitchcock 22271 (US); Oroya, Hitchcock 22184 (US); Hacienda Atocsaico, near lunin, Hitchcock 22199 (US). Distribution and habitat. Trisetum oreophilum var. oreophilum is found in Ecuador, Peru, and Bolivia, along the Cordillera de Los Andes, between 2900 and 4600 m. This species occurs in moist grasslands and ravines. Phenology. Flowering between November and May. Comments. Renvoize (1998) considers Trisetum oreophilum a synonym of T. spicatum. As LouisMarie (1928, 1929) correctly pointed out, T oreophilum is a very homogeneous taxon, apparently more closely allied to T rosei Scribn. & Merr. (both species share pubescent lemmas and a caespitose growth habit) than to T. spicatum. Trisetum oreophilum can be distinguished from T. rosei by having scabrous awns (plumose in T. rosei), smaller spikelets (3.5•4.5 mm long vs. 5•7 mm long in T. rosei), subequal glumes (unequal in T rosei), and glumes and paleas with scabrous keels (vs. ciliate in T rosei). Trisetum rosei grows in Guatemala and Mexico (Espejo-Serna et al., 2000; Finot et al., 2004). Additional specimens studied. BOLIVIA. La Paz: Region Andina, Buehtien 6468 (US); Andean Region, Buchtien 8839 (S); vicinity of La Paz, Mandon 1309 (CONC), 1857 (BAA); Murillo, Valle de Palca, Huasicapampa, Asplund 1028, 2057 (S); Chijini, Asplund 2010 (S); Challapampa, Asplund 3798 (S); Omasuyos, Isla del Sol, Yumani, Asplund 3585 (S); Murillo, La Paz Calacoto 18 km hacia Collana, Beck 13788 (SI); Cerro Calvario, Parodi 10127; 10122 (BAA). ECUADOR. Azuay: EN. Cajas, NW Cuenca along Rio Miguir, Peterson, Annable & Poston 8866 (US). Bolivar: Hacienda Talahua, Penland 584 & Summers (F). Chimborazo: 15.5 km SE of main square of Chambo and SE of Rio Bamba on rd. to Alao, Peterson, 555 14b. Trisetum oreophilum var. jomistonii Louis-Marie, Rhodora 30: 237. 1929. TYPE: Argentina. San Juan: Andes of NW San Juan, Arroyo Tambillos, 4300 m, 10 Jan. 1926, /. M. Johnston 6097 (holotype, GH not seen; isotypes, BAA-3409!, SGO-59037!, SI fragm. ex US!, US-1297379!). Perennial, caespitose; culms 10•20 cm tall, glabrous or pubescent below the inflorescence; node 1. Leaf sheaths pilose; upper sheath inflated, striate: liguie 0.5•1 mm long, truncate, dentate-ciliate; blades flat, pilose; lower blades 3•8 cm X 1•1.5 mm, flat to conduplicate; upper blade 1•5 cm X ca. 1 mm, conduplicate. Panicles 2^1.5 X 0.8•1.5 cm, oval, spiciform, dense, interrupted at the base, green and purpie; rachis giabrous or pubescent. Spikelets 3.5^1.5 mm long, 2- or 3-flowered, subsessile or with pedicels up to 3 mm long, scabrous; rachilla ca. 1 mm long, densely pubescent, with 556 Annals of the Missouri Botanical Garden trichomes up to 1.5 mm long; glumes subequal, shorter than the florets or the second glume as long as the florets; keel scabrous; apex acute; first glume 2.7^ X 0.4-0.9 mm, 1- or 3-nerved, the lateral nerves, if present, very short; second glume 3.3• 4.7 X 0.5-1.2 mm, 3-nerved; florets 3.5-4 X ca. 0.7 mm; lemmas awned dorsally, purplish near the apex; apex biaristulate, pubescent; awn 2•3.5 mm long, somewhat twisted at the base, geniculate or curved, scabrous; callus obtuse, with trichomes ca. 0.5 mm long; paleas 2.2•3.5 mm long, shorter or about as long as the lemma, hyaline, 2-nerved, the nerves scabrous; apex bidentate; anthers ca. 1.1 mm long; lodicules ca. 0.5 mm long, bilobed at the apex, hyaline. Caryopses 1.6•2 X 0.3•0.5 mm, glabrous or with 4 to 6 shining curved trichomes at the apex; endosperm liquid. brada Cantarito, entre Quebrada Marancel y Portezuelo de Cantarito, Marticorena et al. 83462-B (CONC); Km 42 Rio del Estrecho, Arancio, Squeo & Leon 94250 (CONC, USL); Quebrada Los Barriales, Arancio, Squeo & Leon 94043 (USL). IV Region: Choapa, Cordillera de Illapel, Caletdn Blanco, Jiles 4246 (CONC); Cordillera de Combarbala, Ha. Ramadilla, Jiles 4801 (CONC); Cajon de Los Pelambres, Teillier 1536 (CONC); Elqui, Cordillera Dona Ana, Quebrada del Negro, Arancio 92129 (CONC, USL); Cordillera de Ovalle, San Miguel-Los Pingos, Jiles 5889 (CONC); Cordillera de Ovalle, Los Pingos, Jiles 5888 (CONC); Cordillera de Dona Ana, 23 Mar. 1994, Arancio s.n. (USL); Cordillera de Dona Ana, Arancio 93021 (USL); Limari, Cordillera de Ovalle, Punta de Huana-Rio Molles, Jiles 4136 (CONC); Dona Rosa, nacimiento de Quebrada Larga, Jiles 2952 (CONC); Vegas San Miguel, Jiles 3649 (CONC, SGO). Region Metropolitana: Santiago, Paso de las Nieves Negras, Gunckel 20465a (CONC); Cajon del Maipo, Gunckel 20295 (US). Distribution and habitat. Trisetum oreophilum var. johnstonii is found in northern Chile and Argentina (27°S•33°50'S) along the Cordillera de Los Andes, between 2900 and 4600 m. Phenology. Flowering between December and February. Comments. Finot (2003) cited by error Johnston 6079 instead of Johnston 6097 as the type of T. oreophilum var. johnstonii. This species is frequently confused with T. preslei, from which it differs in having hairy leaves (vs. leaves glabrous in T. preslei), glumes shorter than the florets (vs. glumes equaling or exceeding the florets in T. preslei), first glume 1.1•^ mm long (vs. first glume 4.5• 6 mm long in T. preslei), and second glume 3.3•4.7 mm long (vs. second glume (4.5•)5.5•6.5 mm in T. preslei). Additional specimens studied. ARGENTINA. Mendoza: Dpto. San Rafael, Paso Cruz de Piedra, Refugio Peron, Roig 50 (MERL); San Carlos, camino a Laguna Diamante, Boelcke 4132 (BAA); San Carlos, Laguna Diamante, Boelcke et al. 10011, 10039 (BAA); Laguna Diamante, El Paramillo, Boelcke et al. 10061 (BAA); Lujan, Lagunita del Plata, Trombotto & Ahumada 11105 (MERL); Lujan, Cordillera Frontal, Cordon del Plata, Lagunita del Plata, 22 Feb. 1984, Trombotto s.n. (MERL); Las Cuevas, Quebrada Benjamin Matienzo, Perez Moreau 144 (BA, pro parte); Cordillera del Portillo de la Llareta entre el Paso del Portillo y la Laguna del Diamante, entre Arroyo de La Cascada y Corrales Negros, Kurtz 11086 (SI). Neuquen: Chos Malal, cajon inferior del Arroyo Turbio (Arroyo Domeyko), Boelcke et al. 11318 (BAA). San Juan: Calingasta, entre Paso Espinacito Sur y Quebrada Honda, 12 Feb. 1950, Perez-Moreau & Perrone s.n. (BA); Cordillera de Colanguil, quebrada del Salto, Perez-Moreau 30•258 (BA, BAA); Espinacito, Los Frias, Rio Las Lenas, Roig 11950 (BAA); alta Cordillera de San Juan, Laguna Pachon, Koptaluti & Gomez 5816 (SI); Dpto. Iglesia, Mina Fierro Nuevo, Quebrada de los Chilenos, 26 Feb. 1950, Perrone s.n. (BA). CHILE. Ill Region: Copiapo, Laguna del Negro Francisco, Munoz 3981 (SGO); Huasco, Que- 15. Trisetiun phleoides (d'Urv.) Kunth, Revis. Gramin. 1: 101. 1829. Basionym: Avena phleoides d'Urv., Fl. lies Malouin. 30(19). 1825. Trisetum subspicatum var. phleoides (d'Urv.) Hack., Svenska Exped. Magell. 3(5): 222. 1900. Trisetum spicatum subsp. phleoides (d'Urv.) Macloskie, Rep. Princeton Univ. Exp. Patagonia, Botany 8: 206. 1904. TYPE: South America. Falkland Islands [Islas Malvinas], Soledad, d'Urville 3: (holotype, P not found; isotypes, BAA-3411!, US-fragm. ex P!). Trisetum hirsutum Phil., Anales Univ. Chile 46(43): 565. 1873, non (Gaudin) Schrad., Linnaea 12: 443. 1838. Trisetum spicatum (L.) K. Richt. var. hirsutum LouisMarie, Rhodora 30: 239. 1929. TYPE: Chile, "Estrecho de Magallanes, de los alrededores de la colonia chilena," R. A. Philippi s.n. (holotype, SGO-PHIL-235!; isotypes, SGO-63600!, 37057!, US-81775!, US fragm. ex SGO-PHIL-235 & photo!, US photo ex SGO-37057!). Perennial, with short rhizomes; culms 5•24 cm tall, erect, densely pilose below the panicle; trichomes 0.5•0.7 mm long, antrorse below the panicle, then retrorse below. Leaf sheaths pilose; ligule 1.5•2 mm long, truncate, ciliate, dorsally pubescent; blades flat or conduplicate, pilose, ciliate on margins near the ligule. Panicles 2•6 X 0.5•1.5 cm, spiciform, dense, green-yellowish, sometimes tinged with purple, sometimes included in the upper sheath. Spikelets ca. 6.5 mm long, 2- to 4flowered; rachilla ca. 0.8 mm long, pubescent, the trichomes up to 2 mm long; glumes subequal, longer than the florets, rarely the first glume a little shorter than the spikelet; keels ciliate; apex short aristulate; first glume 4•6.5 X 0.5•0.6 mm, 1nerved; second glume 5•6.5 X ca. 0.7 mm, 3nerved; lemmas dorsally awned, glabrous; apex 2aristulate, the awns 0.5•1 mm long; awn ca. 6 mm long, curved, not strongly twisted or geniculate, Volume 92, Number 4 2005 Finot et al. Trisetum in South America scabrous, borne on the upper 1/3; callus with short trichomes, the trichomes ca. 0.2 mm long; florets ca. 4 mm long; paleas ca. 3.5 mm long, shorter than the lemma, 2-nerved, the nerves scabrous; apex bidentate; anthers ca. 1.2 mm long; ovary glabrous. Caryopses not seen. bosque al salto del Rio Grande, 30 Jan. 1912, Hicken s.n. (SI); Isla de Los Estados, Puerto Cook, Mar. 1882, Spegazzini s.n. (LP). CHILE. XII Region: Patagonia Occ., 5 Jan. 1897, Dusen s.n. (S); Magallanes, Anderson 387 (S); Punta Arenas, cerros de canchas de sky, Ricardi & Matthei 316 (CONC); cerros de canchas de sky, 120 m, Pfister & Ricardi s.n. (CONC); Punta Arenas, Rio de las Minas, Barrientos 224 (CONC); Tierra del Fuego, 1769, Banks & Solander s.n. (S); Tierra del Fuego, sector Vicuna, Lote 12, Forestal Trillium, Pisano, Henriquez & Dominguez 7566 (CONC); Isla Navarino, Tsujii 172 (CONC); Isla Navarino, Mont au dessus du Port Williams, Nov. 1958, de La Rile s.n. (P); Fuegia Orientalis, Feb. 1879, P. Ortega s.n. (SGO); Prov. Tierra del Fuego, Rio Hondo, Pisano 2451 (CONC). Distribution and habitat. Trisetum phleoides is endemic to southern Argentina and Chile, occurring between 50°S and 55°S latitudes and between 15 and 1000 m. Phenology. Flowering between November and March. Illustrations. Nicora (1978: 247, fig. 161A-D). Comments. Trisetum phleoides is perhaps related to T spicatum. Both species have spiciform panicles and culms densely pilose below the inflorescence. Trisetum phleoides was placed as a variety or subspecies of T spicatum by Macloskie (1904), Louis-Marie (1928, 1929), and Hulten (1959). Nicora (1978) accepted this taxon at the species level. It is evident from an annotation label written in 1935 by L. Parodi on the type of Avena phleoides at BAA that he considered T phleoides a good species and closely related to T spicatum: "Es una buena especie limitada a las Malvinas y Region Fueguina, afm a T. subspicatum." Trisetum phleoides differs by having pilose blades (vs. glabrous or puberulent in T. spicatum), dorsally pubescent ligules (vs. glabrous in T spicatum), glumes that are ciliate on the keels (vs. scabrous in T spicatum) and longer than the florets (vs. glumes shorter than the florets in T. spicatum). Additional specimens studied. ARGENTINA. Santa Cruz: Rio Fosiles, 1905, Dusen s.n. (S); Giier Aike, Estancia Las Viscachas, Pan de Azucar, Arroyo, Boelcke, Gomez, Moore & Romanczuk 491 (SI); Guer Aike, curso superior del Rio Turbio, entre Estancia La Primavera y puesto Tres Marias, Roig et al. 444 (SI); P.N. Perito Moreno, Rio Chico, Villamil 8396 (CONC). Tierra del Fuego: Ushuaia, cerro Martral, 5 Feb. 1986, Roig J. s.n. (MERL); Valle de Olavaia, Alboff 1039 (SI); Rio Fuego, 9 Jan. 1919, Pico s.n. (BAA); Ushuaia, Mar 1902, Skottsberg s.n. (S); Ushuaia, Skottsberg 236 (S), Feb. 1896, Alboff s.n. (LP); Hito XIX, 6 Feb. 1942, Castellanos s.n. (BAA); alrededores de Ushuaia, Hunziker 8200 (BAA); Ushuaia, canal Beagle, Punta Segunda, Grondona 7340 (BAA); Isla de los Estados, Puerto Abrigado, Castellanos 1937 (BA); Salto del Rio Grande, 30 Jan. 1912, Hicken s.n. (CONC); Islas Malvinas, Skottsberg 69 (S); Ushuaia, Monte Olivia, orillas del Rio Olivia, Ferraro, Messuti & Vobis 4676 (CTES); Ushuaia, Rio Pipo, Camping Mun. Monsefior Aleman, 11 Mar. 1995, Ferraro, Messuti & Vobis 4703 (CTES); Ushuaia, peninsula detras del aerodromo (La Mision), Luti 1640 (CORD); Ushuaia, cerros proximos al monte Olivia, Luti 1426 (CORD); Ushuaia, Ruta Nacional J, Puerto Haberton, Fortunato 4823 & Elechosa (CORD); Ushuaia, Pennington 450b (SI); Turbera de Oldenbourg, F. Roig, C. Roig & FA. Roig 14911 (MERL); camino del 557 16. Trisetum preslei (Kunth) E. Desv., in Gay, Fl. Chil. 6: 347. 1854. Basionym: Avena preslei Kunth, Enum. PL 1: 304. 1833. TYPE: Chile. "Hab. in Cordilleris chilensibus," T Haenke s.n. (holotype, PR-198805!; isotypes, BAA3413 fragm.!, LE-TRIN-1933.02!, MO2106485 not seen, US-81803!). Trisetum lasiolepis E. Desv., Fl. Chil. 6: 346. 1854. Syn. nov. TYPE: Chile. C. Gay s.n. (holotype, P!; isotypes. BAA fragm. ex P not seen, CONC fragm. ex P!, GH not seen, P!, SGO photo ex P!, US-91366 fragm. ex P! >- Deschampsia lasiantha Phil., Linnaea 33: 290. 1864. Trisetum preslei var. lasianthum (Phil.) Louis-Marie, Rhodora 30: 238. 1929. TYPE: Chile. Andes de Hurtado, C. Gay s.n. (holotype, SGO-63603!; isotype. SGO-71900 photo!). Trisetum buchtienii Hack., Z. Bot. 54: 290. 1904. Trisetum preslei var. buchtienii (Hack.) Louis-Marie, Rhodora 30: 238. 1929. TYPE: Chile. "Las Calaveras, 3200 m, Uspallata Pass, der chilenischen Hoch-Cordillere," 14 Feb. 1903, 0. Buchtien s.n. (holotype, S!; isotypes, BAA-3361!, US-1099519!). Perennial, caespitose; culms 3•19(•30) cm tall, erect or geniculate at the base, densely pilose or tomentose below the inflorescence, glabrous below; nodes 1 or 2. Leaf sheaths up to 6 cm long, glabrous; upper sheaths usually inflated and striate; ligule 0.5•1.5 mm long, a little longer on the upper leaves, oval, truncate, dentate-laciniate, ciliate; blades glabrous, adaxially scabrous especially toward the base; margins scabrous; lower blades 2• 6 cm X 1•1.5 mm; upper blades 0.5•3 cm long. Panicles 2•7 X 0.5•1.5 cm, contracted, subspiciform or spiciform, densely-flowered, green or purple, exerted or included in the upper sheath; rachis covered by long trichomes or subglabrous; pedicels up to 2 mm long, pubescent. Spikelets 5•6 mm long, (l)2(3)-flowered; rachilla 1•1.5 mm long, with trichomes 1.5•2 mm long; glumes subequal, equaling or more frequently longer than the florets, delicate, translucid, with wide hyaline margins; keel scabrous; apex acute or aristulate; first glume 4•6 558 Annals of the Missouri Botanical Garden X 0.4•0.7 mm, linear-lanceolate, equaling or a little longer than the spikelet, a little narrower than the second glume, 1-nerved; second glume (4.5•) 5.5-6.6 X (0.6•)0.8•1.1 mm, usually longer than the spikelet, 3-nerved; florets 4.2•5.5 X ca. 0.7 mm; lemmas dorsally awned, green-yelfowish, somewhat purplish toward the apex, dorsally pubescent, trichomes long, soft, up to 1 mm long; margins hyaline; apex 2-aristulate; awn 2.5^1.5 mm long, nearly as long as the lemma, borne on the upper 1/3, divaricate, geniculate and weakly twisted or more frequently curved not geniculate or twisted, pilose at the base, scabrous above; callus obtuse, pilose, with trichomes ca. 1 mm long; paleas 3.2•4.5 mm long, a little shorter than the lemma, 2-nerved, the nerves scabrous-ciliate; apex 2dentate; lodicules 0.6•1 mm long; apex bilobed, one of the lobes larger; ovary glabrous. Caryopses 2.5•2.7 X ca. 0.5 mm, glabrous; endosperm dry. Additional specimens studied. ARGENTINA. Mendoza: on road above Las Cuevas, below Cristo Redento, H. & B. Mooney 546 (CONC); La Cumbre, Las Cuevas, Dec. 1908, Spegazzini s.n. (LP); Valle del Atuel, cerca Laguna Atuel, Bbcher et al. 1971 (BAA); Atuel Valley, near El Angulo, Bbcher et al. 1894 (BAA); Las Heras, entre Las Cuevas y Cristo Redentor, Ruiz Leal 79 (MERL); Las Heras, entre Las Cuevas y Cristo Redentor, Ruiz Leal 6625 (MERL); Las Heras, Cristo Redentor, Martinez Carretero 1272 (MERL); San Rafael, valle del no Atuel, Boelcke et al. 10226 1/2 (BAA); San Rafael, Antes de llegar Indigeno-Estribaciones Norte Volcan Overo-El Sosneado, Lagiglia 2239 (LP); Agua Amarilla, Volcan Overo, El Sosneado, Lagiglia & DAntoni 1326 (LP); entre Puesto de Ubilla y La Manzanilla, en el arroyo Tordillo, Kurtz 7599 (CORD); Calmuco, Covas 101 (BAA); Malalhue, proximidades de la cresta de la Sierra Azul, Mendez & Willoud c-320-7196 (MERL). Neuquen: Chos Malal, Cajon del Arroyo del Cruce, faldeo S del Domeyko, Boelcke et al. 11285 (BAA); Lacar, Cerro Repollo, Estancia Meliquina, Ritgolo de Agrasar & Agrasar 5846 (BAA); Minas, Lagunas Epu-Lauquen, Puesto de Gendarmena, Boelcke et al. 11033 (BAA); Pino Hachado, Parodi 3196 (US); Chos Malal, a 34 km de Tricao Malal camino a Mina de Azufre, Boelcke et al. 11674 (SI). Rio Negro: El Bolson, Cerro Piltriquitrdn, Cabrera et al. 23110 (LP); Parque Nacional Nahuel Huapi, Cerro Lopez, Boelcke 1972 (BAA). CHILE. IV Region: Coquimbo, Los Molles, Ovalle, Zoellner 5859 (CONC). V Region: Los Andes, Laguna Castro, Penaloza et al. 91126, 91121 (CONC); entre Laguna Las Truchas y Laguna de la Turquesa, Arroyo, Maldonado & Henriquez 91156, 91158 (CONC); Laguna Castro, Penaloza et al. 91122 (CONC); Aconcagua, Portillo, Laguna del Inca, Sparre 1678 (S). Region Metropolitana: Cordillera de Santiago, Mar. 1899, Reielie s.n. (SCO); Parque Nacional El Morado, Cordillera de Los Andes frente a Santiago, Teillier et al. 2548 (SGO); San lose de Maipo, Cajon del Rio Morales, Saavedra & Pauchard 6 (CONC, SGO); Cordillera, Lo Valdes, Feb. 1950, Gunckel s.n. (CONC); Cajon del Maipo, Gunckel 20297 (US); Cajon del Maipo, Hito Paso Internacional Maipo, Villagrdn et al. 8462 (SGO); Maipo, Joseph 2948 (US); Prov. Santiago, Camino de Santiago a Mina La Disputada, 2 km antes de Perez Caldera, Marticorena & Matthei 664 (CONC); Cajon del Yeso, Termas El Plomo, M. Munc-z et al. 3491 (SGO); Laguna Negra, Lechler 2948 (US). VIII Region: Nuble, Termas de Chilian, Cabrera 3662 (LP), Feb. 1947, Castillo s.n. (CONC, US); Bafios de Chilian, Ian. 1877, F. Philippi s.n. (SGO), Ian. 1878, F. Philippi s.n. (LP). Distribution and habitat. Trisetum preslei is endemic to Chile and Argentina. In Chile T. preslei is found along the Cordillera de Los Andes between 2200 and 3700 m, and in Argentina it occurs between 33°S and 42°S at 1800-4000 m. Phenology. Flowering between December and February. Illustrations. Nicora (1978: 262, fig. 169A-C). Comments. A note by Desvaux (1854) accompanying the original description of T. lasiolepis indicates a close relationship of this species with T. preslei. In Desvaux's opinion T. lasiolepis could be considered as a variety of T. preslei. Trisetum lasiolepis was recognized as a good species by LouisMarie (1928, 1929) and by Nicora (1978). Nicora (1978) distinguished T. lasiolepis from T. preslei as being taller (plants 20^10 cm tall in T. lasiolepis vs. 10•20 cm tall in T. preslei), culms glabrous or pilose just below the panicle (vs. retrorse tomentose in T. preslei), panicles somewhat loose, with rachis pubescent to almost glabrous (vs. panicle dense, with rachis densely tomentose in T. preslei), and keels of glumes and palea scabrous (vs. keels of glumes and palea normally ciliate in T. preslei). The type of T. lasiolepis at P has culms 20•30 cm tall, narrow subspiciform panicles 5•7 cm long and less than 1 cm wide, glabrous blades (lower blades 4• 5 cm long; upper blades 1•2 cm long), 2-flowered spikelets 5•6 mm long, and glumes equaling the florets or the second glume longer than the florets (first glume 4 X 0.4 mm; second glume 6 X 0.8 mm). Since the morphological characters taken from the type of T. lasiolepis fall within the range given for T. preslei, we consider this a new synonym of the latter species. 17. Trisetum pyramidatum Louis-Marie ex Finot, sp. nov. TYPE: Chile. Punta Arenas, Lena Dura, 28 Jan. 1946, M. Barros 5706 (holotype, US-1869901!). Figure 3. Planta perennis, rhizomatosa, 37•50 cm alta; culmi pilosi, inferiores glabri; folia glabra; laminae planae; ligula 2•3 mm longa, glabra; panicula pyramidata, 7•11 X 2•3 cm; spicula 6•6.5 mm longa, 2- v. 3-flora; glumae subaequales vel inaequales, spiculam aequantes, ad apicem aristulatae; gluma I: (5•)5.5•7.5 X 0.8•1 mm, 1-nervia; gluma II: 6.5•9 X 1.1•1.3 mm, 3-nervia, quam lemma sua minores vel excedentes; lemma ad dorsum scabra, ad terciam superiorem aristata; arista geniculata vel recurvata, scabra, 6•7 mm longa; callus brevi-pilosus; rhachilla pilosa 1 mm longa; palea quam lemma minor, binervia, ad Volume 92, Number 4 2005 Finot et al. Trisetum in South America 559 Figure 3. Trisetum pyramidatum. •A. Habit. •B. Sheath, ligule, and portion of the blade. •C. Spikelet. •D. Florets. •E. Palea, dorsal view. •F. Palea, ventral view. •G. Rachilla. •H. Lodicules. •I. Stamen. •J. Caryopsis. Barros 5706 (holotype, US). 560 apicem bisetulosa; stamina 3, antheris 0.5 mm longis; lodiculae 0.8 mm longae, ad apicem bilobatae; ovarium glabrum; caryopsis 2.5 mm longa, glabra; endospermium granulosum. Perennial, with short rhizomes; culms 37•50 cm tall, pubescent below the panicle, with trichomes first antrorse, then retrorse; nodes 2, glabrous. Leaf sheaths glabrous, pubescent when young; ligule 2• 3 mm long, membranous, with rounded, dentate and ciliate margin; dorsally glabrous; blades 4•7 cm X 2•2.3 mm, flat, glabrous; lower blades 6•7 cm X 2•2.3 mm; upper blades 4•5 cm long, conduplicate toward the apex. Panicles 7•11 X 2•3 cm, contracted, pyramidal, yellowish-green and tinged with purple, bright; branches in trimerous whorls, the basal branches up to 3 mm long. Spikelets 6•6.5 mm long, 2- or 3-flowered; glumes subisomorphic or dissimilar, equaling or longer than the spikelet or at least the second glume longer than the spikelet, wide, lanceolate, adaxially purplish; margin hyaline; apex aristulate, hyaline; first glume (5•)5.5•7.5 X 0.8•1 mm, 1-nerved; second glume 6.5•9 X 1.1•1.3 mm, 3-nerved; florets 6.5• 7 mm long; lemmas dorsally awned, scabrous; apex hyaline, biaristate; awn 6•7 mm long, borne on the upper 1/3, geniculate or only diversely curved, not twisted, scabrous, purplish; callus obtuse, with very short trichomes, the trichomes 0.2•0.3 mm long; rachilla ca. 1 mm long, pubescent, with trichomes ca. 1.5 mm long; paleas 4•5 mm long, shorter than the lemma, 2-setulate at the apex, 2-nerved, the nerves scabrous; anthers ca. 0.5 mm long; lodicules ca. 0.8 mm long, 2-lobed, the lobes acute; ovary glabrous. Caryopses ca. 2.5 mm long, glabrous; endosperm dry, granular. Distribution and habitat. Trisetum pyramidatum is endemic to Chile and Argentina, between 40°S and 53°S at low elevations less than 200 m. Phenology. Flowering between December and March. Comments. Trisetum pyramidatum is morphologically similar to T. phleoides. This new species can be distinguished from T. phleoides by having wider, pyramidal-shaped panicles, (vs. spiciform in T. phleoides) and by having glabrous blades (vs. densely pubescent in T. phleoides). Paratypes. ARGENTINA. Rio Negro: Nord Patagonia, San Carlos de Bariloche, Feb. 1905, Buchtien s.n. (US). CHILE. XI Region: Aisen, Quitralco, 15 Dec. 1947, Behn s.n. (CONC). XII Region: Punta Arenas, Lena Dura, Barros 5709 (US); Punta Arenas y Rio de la Mina, 1 Mar. 1917, Bonarelli 185 (SI); Rio Lena Dura, al sur de Punta Arenas, Bonarelli 181 (SI); Rio Tres Brazos, al sur de Punta Arenas, Bonarelli 182 (SI). 18a. Trisetum spicatum (L.) K. Richt. var. spi- Annals of the Missouri Botanical Garden catum, PL Eur. 1: 59. 1890. Basionym: Aira spicata L. Sp. PL 1: 64. 1753. Aira subspicata L., Syst. Nat. ed. 10. 2: 873. 1759, nom. illeg. superfl. Avena airoides Koeler, Descr. Gram. 298, 1802, nom. illeg. superfl. Trisetum subspicatum (L.) P. Beauv., Ess. Agrostogr. 88, 149. 1812, nom. illeg. superfl. Trisetaria airoides (Koeler) Baumg., Enum. Stirp. Transsilv. 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.) Reichb., Fl. Germ. Excurs. 49. 1830, nom. illeg. superfl. Koeleria spicata Reichb. ex Willk. & Lange, Prodr. Fl. Hispan. 1: 72. 1861, nom. inval. Trisetaria spicata (L) Paunero, Anales Jard. Bot. Madrid 9: 516. 1959. TYPE: Sweden. Lapland: 1732, Linnaeus s.n. (lectotype, LINN 85.7! designated (as holotype) by Edgar, New Zealand J. Bot. 36: 556. 1998; isotype, S!). Trisetum albidum Sodiro, Revista Col. Nac. Vicente Rocafuerte 12: 84, 86. 1930. TYPE: Ecuador. Pichincha: crece en los potreros interandinos, Quito, Chillogallo, Cotocollao, and Pifo, Sodiro s.n. (holotype. not located; isotype, US-1163185 fragm.!). Trisetum andinum Phil., Linnaea 29: 93. 1858. horn, illeg. non Benth. 1847. TYPE: Chile. In andibus prope Antuco invenit C. Gay Herb. Chil. 210 (holotype. SGO-PHIL-241!; isotypes, SGO photo!, US fragm. ex SGO-PHIL-241! & photo!). Trisetum spicatum subsp. bolivianum Hulten, Svensk Bot. Tidskr. 53: 224. 1959. TYPE: Bolivia. La Paz: Murillo, at the railway station La Cumbre, ca. 4700 m. 28 Apr. 1921, Buchtien 8839 (holotype, S!). Perennial, caespitose or with short rhizomes; culms 9•60 cm tall, erect, tomentose to densely pubescent below the panicle, with trichomes antrorse below the panicle, then retrorse below; nodes 1 or 2. Leaf sheaths 1•3(•6) cm long, glabrous; ligule ca. 1 mm long, finely denticulate; blades 1• 5 cm X 1•1.5 mm, flat or conduplicate toward the apex, glabrous or rarely pubescent or scabrous, sometimes ciliate on margins. Panicles 2.5•7(•10) X 0.5•1.5(•2) cm, spiciform, gold-purplish to brown-purple, bright, usually interrupted at the base; rachis pubescent; pedicels pubescent. Spikelets 4.5•6 mm long, 2-flowered; rachilla 0.8•1 mm long, pubescent, the trichomes 0.5•1 mm long; glumes subequal, shorter than the florets, as long as 3/4 to 4/5 of the spikelet, subequal or the first a little shorter and narrower than the second glume; sometimes, the second glumes equal or a little longer than the florets, scabrous or less frequently ciliate on the keel; first glume 3.7•5 X 0.5•1 mm, lanceolate, 1- to 3-nerved; second glume 4.5•6 X 0.5-1.3 mm, 3-nerved; florets 3.8-5 X 0.7-0.8 Volume 92, Number 4 2005 Finot et al. Trisetum in South America mm, the second floret 4•4.5 mm long; lemmas dorsally awned, glabrous, scabrous, purplish toward the base, stramineous toward the apex; margin hyaline; apex 2-aristulate; awn 3.5•5 mm long, borne on the upper 1/3 or 1/4, geniculate or merely curved, sometimes twisted, scabrous, purple; callus obtuse, with trichomes 0.3•0.5 mm long; paleas 3• 4 mm long, shorter or a little longer than the lemma, hyaline, 2-nerved, the nerves scabrous; anthers 0.5•1 mm long; lodicules ca. 0.6 mm long, hyaline; apex 2-lobed; ovary glabrous. Caryopses 2•2.8 X ca. 0.6 mm, glabrous; endosperm liquid. er, second ca. 4X2 mm) glabrous; lemmas (ca. 4 mm long) awned dorsally on the upper 1/4; awn (ca. 4 mm long) reflexed; palea equal or subequal to the lemma; rachilla densely villous; anthers 0.6• 1 mm long." This description corresponds well with the isotype of Aira spicata L., kept in Stockholm Distribution and habitat. Cosmopolitan, probably native to Europe, introduced in America (Tovar, 1993). Trisetum spicatum has a circumpolar distribution (Hulten, 1959; Clebsch, 1960). In the Southern Hemisphere it is found in South America, Australia, and New Zealand (Clebsch, 1960). In North America it grows in Canada, United States, Greenland, and Mexico. In South America T. spicatum extends from Colombia to southern Chile and Argentina (5^1700 m) and reaches its southern limit in Tierra del Fuego at 55°S, where it is found in bogs associated with Empetrum L. (Empetraceae) and Azorella Lam. (Apiaceae). Pohl and Davidse (1994) reported T. spicatum for Guatemala, Tovar (1993) found this species in central and southern Peru along the Cordillera de Los Andes between 3900 and 4700 m, and Renvoize (1998) recorded this species for Bolivia (La Paz, Cochabamba, Oruro). In Argentina, Zuloaga et al. (1994) reported T. spicatum as occurring in Neuquen, Rio Negro, Santa Cruz, and Tierra del Fuego. Phenology. Flowering between October and March. Comments. Trisetum spicatum has been treated as a complex including a vast number of subspecies and/or varieties (Louis-Marie, 1928, 1929; Hulten, 1959). Louis-Marie recognized 14 varieties, and most of these were not recognized as valid entities by later authors. Hulten (1959) divided the species into 22 infraspecific taxa, including 14 subspecies and 8 varieties. Based on a multivariate analysis of 33 morphological characters, Randall and Hilu (1986) could not differentiate any clear-cut forms or infraspecific taxa within T spicatum. Louis-Marie (1928, 1929) transcribed the description made by Scheuchzer in 1719 and cited by Linnaeus (1753) in his diagnosis of Aira spicata: "The plant (17•30 cm high) is described as having glabrous, striate blades; ligule (ca. 1 mm long) obtuse; culms densely tomentose; panicles spike-like (ca. 1.5 X 0.7•1 cm), purplish, shining; spikelets (ca. 4 mm long) 2flowered; glumes unequal (first shorter and narrow- 561 (S). The type specimen of Trisetum albidum (US1163185) has somewhat lax panicles and shorter spikelets (ca. 4 mm long) than in typical T. spicatum (4.5•6 mm long). The glumes are, in consequence, shorter (first glume 3•3.2 X 0.4•0.5 mm; second glume 3.6^1 X 0.8 mm). Since these measurements fall within the range of the original description given for T. spicatum in the last paragraph, we choose to relegate this form to synonymy, even though earlier VLF tentatively accepted T. albidum (Finot, 2003). Chromosomes. 2n = 28, 42 (Tateoka, 1978). Illustrations. Hitchcock and Chase (1950: 290, fig. 390); Hulten (1959: 207, fig. 1); Nicora (1978: 247: 169A-D). Additional specimens studied. ARGENTINA. Chubut: Lago La Berta, Nicora 9603 (SI); Lago Vinter, Roig et al. 14266 (SI, MERL); Lago Vinter, Nicora 10235, 10284, 10344, 10348 (SI), Nicora 9485 (CONC); Languineo, Lago El Guacho, Nicora 9519 (CONC); Lago Cuatro. Nicora 9344 (CONC); Rio Corcovado, Illin 169 (BAF); Lago La Para, Nicora 10081 (SI); Valle Huemules, Soriano 3184 (BAA). Cordoba: San Alberto, Sierra Grande, Pampa de Achala, Ea. La Trinidad, 25 Jan. 1984, Cabido s.n. (CORD). Neuquen: Los Lagos, Arroyo Pantojo, cascada Santa Ana, Rugolo de Agrasar 1093 (SI); Dpto. Minas, a 21 km de Las Ovejas, camino a las lagunas Epu-Lauquen, arroyo de Las Bandurrias, Boelcke et al. 10801 (BAA); Cerro Malo, Schajowskoy 55-a (BA); Lago Nahuel Huapi, Paso de las Nubes, Cabrera 5928 (LP); Pino Hachado, Puesto Gendarmeria, junto al arroyo. Valla et al. 3065 (CTES); Andacollo, Arroyo Guaraco, Cabrera 11143 (LP). Rio Negro: PN. Nahuel Huapi, paso de Las Nubes, Boelcke & Correa 5590 (BAA); Valle del Rio Alerce, Boelcke & Correa 5631 (BAA); Cerro Tronador, Mallin Chileno, Boelcke & Correa 5775 (BAA); Cerro Tronador. Job 2431 (LP); Lago Frias, Feb. 1943, Soriano s.n. (BAA). Santa Cruz: Spegazzini s.n. (LP); Lago Viedma, 19 Feb. 1905, Dusen s.n. (S); Lago Frias, Roquero 31 (BA); Cerro San Lorenzo, Roquero 297 (BA); Sierra Colorada, Dimitri & Correa Luna 119 (BA); PN. Los Glaciares, Lago Argentino, Ea. Cristina, Roquero 434 (BA); Rio Chico, Dec. 1897, Ameghino s.n. (LP); Lago Argentino, Sierra de Buenos Aires, Roquero 169 (BA); Lago Argentino, James 189 (SI); La Vizchacha, Burmeister 3 (SI); Guer Aike, Ea. Sofia, Secc. Cuadrado, 5 km S de Estancia Punta del Monte, TBPA 3117 (SI); Ea. La Verdadera Argentina, Cerro de La Virgen, Arroyo et al. 215 (TBPA 2260) (SI); Ea. Stag River, 10 km N del casco sobre el no de los Venados, TBPA 3140 (SI). Tierra del Fuego: Isla de los Estados, Puerto San Juan, 28 Dec. 1933, Castellanos s.n. (BAA); Rio Grande, lado camino, entre Punta Centolla y Cabo Viamonte, 25 Dec. 1970, Panza s.n. (BAA). BOLIVIA. La Paz: Nor Yungas, 8 mi. E of Pass on road to the Yungas 562 Annals of the Missouri Botanical Garden (Unduavi), Peterson, Soreng & Laegaard 13173 (US); Murillo, Pass at the head of the Valle del Zongo and lower slopes of Nevado Hayna Potosi, Solomon 13221 (SI ex MO); 2 km N of Millumi, Lara & Parker 41J (F). CHILE. V Region: Valparaiso, near summit La Campana, 10 mi. E of El Granizo, Eyerdam 10047 (F, SGO). Region Metropolitana: Santiago, Quebrada El Yeso, Araya 16 (CONC). VII Region: Curico, Andes de Curico, Vidal 233, 234 (US, SGO). VIII Region: Nuble, Termas de Chilian, Jaffuel 1806-a (SGO); Concepcion, Tomeco, Barros 3982 (LP); Parque Nacional Laguna del Laja, Los Barros, sector Aduana, Finot & Baeza 12 (CONC); entre Chacay y canchas de sky, Finot & Baeza 2071 (CONC). IX Region: Malleco, Fundo Solano, Los Alpes, Cordillera de Nahuelbuta, Eyerdam 10257 (F). X Region: Valdivia, Volcan Quetrupillan, fundo Trafun, Schlegel 7523 (CONC); Valdivia, Panguipulli, lago Rifhhue, lado norte, Montero 9557 (CONC); Osorno, Antillanca, Sparre & Constance 10778 (CONC); Antillanca, Schlegel 7322 (CONC); Llanquihue, Volcan Yates, Werdermann 655 (SI); Cerro Vichadero, Casa Pangue, 14 Jan. 1953, Pfister s.n. (CONC). XI Region: Aysen, Munoz s.n. (CONC); Puyuhuapi, Cerro Tesoro, Schwabe 69 (CONC); Coihaique, Barros 5719 (US); Estero Cofre, Vogel 545 (CONC). XII Region: Parque Nacional Torres del Paine, Cerro Diente, Arroyo & Squeo 860055-a (CONC); Isla en el lago Grey, Rugolo de Agrasar 1176 (CONC); Magallanes, Rubens, 10 Jan. 1952, Pfister & Ricardi s.n. (CONC); Tierra del Fuego, Caleta Josefina, Ricardi & Matthei 177 (CONC). COLOMBIA. Caldas: Cordillera Central, vertiente occidental, paramos del Nevado del Ruiz, Cuatrecasas 9280 (F). ECUADOR. Chimborazo: Fagerlind & Wibon 833 (S). Cotopaxi: 19 km east of Pilalo, Peterson, Annable & Poston 8765 (US); Carchi, Hacienda La Esperanza, El Voladero, paramo El Angel, Ddvalos 27 (US). PERU. Junin: Reserva Nacional de Junin, Ondores, Pettersson 15 (S). chomes; upper internodes 12•28 cm long; nodes 2 or 3, glabrous or pilose. Leaf sheaths shorter or a little longer than the internodes, glabrous to sparsely pilose; ligules 1.5•3 mm long, glabrous or pilose dorsally, dentate, ciliate; blades 5.0•15 cm X 1• 1.5 mm, flat or conduplicate, pilose; margins ciliate; upper blades 1•3.5 cm long. Panicles 4•10 X 0.8•2 cm, spiciform, green and purplish, usually interrupted at the base, shining; rachis glabrous or pilose; pedicels scabrous or with a few short trichomes. Spikelets 6•8 mm long, 2- or 3-flowered; rachilla 1•1.5 mm long, pilose, the trichomes 0.8• 1 mm long; glumes unequal, acute or arrstulate; keel scabrous-ciliate; first glume 4.5•7 X 0.2•0.6 mm, linear-lanceolate, narrow, shorter and narrower than the second glume, shorter than the florets, 1nerved, attenuate toward the apex; second glume (5•)6•8 X 0.9•1 mm, ovate to ovate-lanceolate, longer than the florets, 3-nerved; lemmas dorsally awned, glabrous, scabrous toward the apex; apex 2aristulate, the awns 0.5•1.5 mm long; awn 6•8 mm long, borne on the upper 1/3, twisted and geniculate to merely curved, scabrous, purple; callus obtuse, with trichomes ca. 0.7 mm long; paleas hyaline, 2-nerved; anthers ca. 1.5 mm long; lodicules ca. 1 mm long; apex bilobed; ovary glabrous. Caryopses glabrous; endosperm liquid. 18b. Trisetum spicatum var. cumingii (Nees ex Steucl.) Finot, comb. nov. Basionym: Koeleria cumingii Nees ex Steucl., Syn. PL Glumac. 1: 294. 1854. Trisetum cumingii (Nees ex Steucl.) Nicora, Fl. Patagonica 3: 250. 1978. TYPE: Chile. Prope Valparaiso, 1831, H. Cuming 460 (lectotype, designated here, BAA-3396 fragm. ex Herb. Nees at B! with illustration of spikelet). Trisetum mollifolium Louis-Marie, Rhodora 30: 218. 1928, nom. nov. Trisetum malacophyllum Phil., Anales Univ. Chile 48: 566. 1873, non Steucl., Syn. PL Glumac. 1: 229. 1854. TYPE: Chile. Tierra del Fuego: Fue igualmente hallada en la vecindad de la colonia chilena, en el Estrecho de Magallanes, Lechler 1195 (holotype, SGO-PHIL-221!; isotypes, BAA-3398!, BAA-3399!; CONC-148141 fragm. & photo ex MI, CONC-148150 fragm. & photo ex B; SI, SGO37056!; US-1126269!, US-868482 fragm. ex PI, US81778 fragm. ex SGO-PHIL-221!, F-881353!). Trisetum cumingii var. santacrucense Nicora, FL Patagonica 3: 250. 1978. TYPE: Argentina. Santa Cruz: Dpto. Lago Argentina, El Calafate, M. N. Correa 2953 (holotype, BAB not seen, isotype, BAA-3370!). Perennial, caespitose; culms 15•50 cm tall, erect, glabrous or with a few short appressed tri- Distribution and habitat. Trisetum spicatum var. cumingii is endemic to Chile and Argentina. In Argentina it is found south of the 40th parallel and in Chile it extends from 33°S (V Region) to southern 50°S, in Ultima Esperanza (XII Region) between 10 and 1100 m. Phenology. Flowering between December and March. Comments. The type of Koeleria cumingii in Berlin-Dahlem (B) was destroyed during WWII (Oberprieler, herb. B, pers. comm.). A fragment of this type was deposited in Buenos Aires (BAA), with a label indicating "Herb. Nees, Valparaiso, Chile, 1831, Cuming." This fragment was also annotated by Prof. Lorenzo Parodi in which he noted "inflorescencia de 10 cm de largo por 1.5 cm de ancho; hoja unica plana, vaina pubescente•Berlin, 1936." Other annotations by Parodi include: "raquis y ramas pilosos; carena glabra; gluma (1)6• 6.5 mm; gluma (2)7•7.7 mm; arista de la lemma 6•7 mm de longitud; lemma aspera?, escabriuscula." This taxon was accepted as a species by Nicora (1978), who comments that the acute glumes are longer than 4 mm and the second glumes are longer than the contiguous florets. We were able to study a large number of specimens of T. spicatum from South America, many of which showed the Volume 92, Number 4 2005 Finot et al. Trisetum in South America characters commonly used to separate T. cumingii Penaloza 870266 (CONC); Las Cumbres, Baguales, Ricardi & Matthei 385 (CONC); Peninsula Munoz Gamero, entrada al lago, Dollenz 11 (CONC); Sector Vicuna, Lote 12, Forestal Trillium, Pisano, Henriquez & Dominguez 7436 (CONC); Manantiales, Magens 3428, 3433 (CONC); PN. Torres del Paine, Rio Grey, Dollenz 1393 (CONC); Lazo, cerca Lago Toro, potrero El Chingue, Munoz 4104, 4144 (SGO); Magallanes, F. Philippi s.n. (SGO). from T. spicatum, specifically the second glume longer than the contiguous floret. Nicora (1978) commented that many specimens collected in Lago Belgrano showed intermediate characteristics between T. spicatum and T. cumingii. Since there are intermediate specimens, we propose to recognize T. cu- 563 mingii as a variety of T. spicatum. Literature Cited Additional specimens studied. ARGENTINA. Chubut: Dpto. Paso de Indios, desvio a 35 km SW Ruta 24. Sa de la Butrera, Estancia La Altura, Corral de Piedra, Arroyo, Mon & Romanczuk 200 (CONC); Lago Puelo, Estancia La Esperanza, picada al S del lago Esperanza, Cusato & Rossoiv 4524 (BAF). Neuquen: Dpto. Los Lagos, Pto. Manzano, Lago Nahuel Huapi, Rugolo de Agrasar 1252 (CONC); PN. Nahuel Huapi, entre Puerto Blest y Lago Frias, Dimitri & Correa 176 (BA); Quetrihue, Diem 458 (BAA). Rio Negro: Cordon Serrucho, El Bolsdn, Roig Junent 13353 (MERL); Lago Fonk, 21 Jan. 1942, Perez-Moreau s.n. (BA); Bariloche, Puerto Blest, Nahuel Huapi, Villamil et al. 2844 (CONC); Bariloche, cerca Rio Nirihuau, Parodi 15582 (BAA); Rio Negro, Bariloche, cerca Rio Nirihuau, Parodi 15560 (BAA). Santa Cruz: Giier Aike, Rio Turbio, Roig et al. 14748 (MERL); Valle Superior del Rio Turbio, faldeo de la Cordillera Chica, Mendez & Ambrosetti s.n. (MERL); Cordillera Chica, Cerro Punta Alta, Ambrosetti & Mendez 29884 (MERL); Valle Superior del Rio Turbio, Meseta Latorre, Ambrosetti & Mendez 29889 (MERL); Valle Superior del Rio Turbio, puesto Tres Marias, Ambrosetti & Mendez 407 (SI); Valle Superior del Rio Turbio, entre Pto. 16 La Primavera y Pto. 3 Marian, Ambrosetti & Mendez 29874 (MERL); Cordillera Chica, Ambrosetti & Mendez 29885 (TBPA) (MERL); Estancia Stag River, faldeos Meseta Latorre, Ambrosetti & Mendez 27457 (MERL); Banado 28 de Noviembre, Ambrosetti & Mendez 29882 (MERL); Valle Superior del Rio Turbio, Ambrosetti & Mendez 29876 (MERL); Lago Argentino, James 247 (SI); Lago Argentino, Sierra Buenos Aires, Roquero 132 (BA); PN. Perito Moreno, Lago Burmeister, Rumboll 120 (BA); Ea. La Carlota, seccion San Elias, Roig, Anchorena, Mendez & Ambrosetti 115 (SI); Estancia Las Viscachas, Cerro Las Viscachas, TBPA 2404 (CONC); Canadon Quitapenas, Correa 4016 (CTES). Tierra del Fuego: Moat, Roig 15589 (MERL), Roig et al. 14912 (MERL); cerca Cerro Mesa, 31 Jan. 1942, Castellanos s.n. (BA); Ushuaia, A. Roig & F. Roig 14980 (MERL, BAA); Rio Grande, Castellanos s.n. (BAA); Lago Fagnano, Monte Huehuepen, Moore & Goodall 398 (CONC). CHILE. VIII Region: Lirquen, alto del camino a Tome, Pfister s.n. (CONC). X Region: Chiloe, Isla Guafo, trayecto desde Caleta Samuel al Faro, Villagrdn & Leiva 7525 (CONC). XII Region: Magallanes, 15 km south of Punta Arenas, Eyerdam, Beetle & Grondona 24133 (US); Punta Arenas, Magallanes 1864/65, Philippi s.n. (SGO); Cerro Castillo, Magens 3197 (CONC); Tierra del Fuego, Ea. Cameron, Ricardi & Matthei 184 (CONC); Tierra del Fuego, Santa Catalina, Ricardi & Matthei 263 (CONC); Laguna Blanca, Riggi 27/1052 (BA); Morro Chico, Magens 3032 (CONC); tJltima Esperanza, Arroyo et al. s.n. (CONC 92112); Ultima Esperanza, Sierra de Los Baguales, Cerro Santa Lucia, Arroyo s.n. (CONC 85168); Sierra de Los Baguales, Cerro Santa Lucia, Arroyo s. n. (CONC 85159); Cordillera del Paine, von Bohlen & Cavieres s.n. (CONC 92312); Cerro Donoso, Sector Rio de Las Chinas, Arroyo, Veloso & Baeza, C. M., G. Kottirsch, J. Espejo & R. Reinoso. 2001. Recuentos cromosomicos en plantas que crecen en Chile. I. Gayana, Bot. 58(2): 133-137. Bolkhovskikh, Z., V. Grif, T. Matvejeva & 0. Zakharyeva. 1969. Trisetum. In: A. Fedorov (editor), Chromosome Numbers of Flowering Plants. Academy of Science of the USSR. V.L. Komarov Bot. Inst. Calderon, C. E. 1978. XL Leptophyllochloa. Pp. 69-71 in E. G. Nicora, Flora Patagonica, part III•Monocotyledoneae: Gramineae. Coleccidn Cientifica del Instituto Nacional de Tecnologia Agropecuaria, Buenos Aires. Calderon de Rzedowski & J. Rzedowski. 2001. Flora fanerogamica del Valle de Mexico. Instituto de Ecologia, A.C., Centro Regional del Bajio, Comision Nacional para el Conocimiento y Uso de la Biodiversidad, Patzcuaro, Michoacan. Chrtek, J. 1965. 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Renvoize, S. A. 1998. Gramineas de Bolivia, i•x. The Royal Botanic Gardens, Kew. Rugolo de Agrasar, Z. E. & E. G. Nicora. 1988. Nuevos taxones para la Argentina y Chile Austral (Gramineae). Bol. Soc. Argent. Bot. 25(3-4): 463^176. Scheuchzer, J. 1719. Operis Agrostographici idea seu graminum, juncorum, cyperorum, cyperoideum, iisque afinium methodus. Tiguri [Zurich]. Steudel, E. G. 1853•1855. Synopsis Plantarum Glumacearum. 2 vols. Part 1. Gramineae. J. B. Metzler, Stuttgart. Swallen, J. B. 1948. New grasses from Honduras, Colombia, Venezuela, Ecuador, Bolivia, and Brazil. Contr. U.S. Natl. Herb. 29: 251-275. Tateoka, T. 1978. Cytotaxonomic conspectus of Japanese Trisetum (Poaceae). Bull. Natl. Sci. Mus. Tokyo, B (Bot.) 4: 1-3. Tovar, 0. 1993. Las gramineas (Poaceae) del Peru. Ruizia 13: 1-480. Tsvelev, N. 1970. De generibus Trisetum Pers. et Koeleria Pers. in URSS. Notulae Systematicae 7: 59•73. . 1983. Grasses of the Soviet Union [Zlaki SSSR]. Botanicheskii Institu im. V L. Komarova and Smithsonian Institution, Amerind Publ., New Delhi. Tucker, G. C. 1996. The genera of Pooideae (Gramineae) in the southeastern United States. Harvard Pap. Bot. 9: 11-90. Valencia, J. I. 1941. Especies criticas de Trisetum que deben pasar al genero Deschampsia. Revista Argent. Agron. 8: 122-130. Veldkamp, J. F. & J. C. Van der Have. 1983. The genus Trisetum (Gramineae) in Malesia and Taiwan. Gard. Bull. Singapore 36: 125-135. Zuloaga, F. 0., E. G. Nicora, Z. E. Rugolo de Agrasar, 0. Morrone, J. Pensiero & A. M. Cialdella. 1994. Catalogo de la familia Poaceae en la Republica Argentina. Monogr. Syst. Bot. Missouri Bot. Gard. 47. APPENDIX 1. ALPHABETICAL INDEX OF ACCEPTED SPECIES AND VARIETIES. NUMBER IN PARENTHESES CORRESPONDS TO THE NUMBER OF THE SPECIES IN THE TAXONOMIC TREATMENT. Trisetum ambiguum Rugolo de Agrasar & Nicora (5) Trisetum andinum Benth. (6) Trisetum barbinode Trin. (7) Trisetum barbinode Trin. var. barbinode (7a) Trisetum barbinode var. hirtiflorum (Hack.) Louis-Marie (7b) Trisetum barbinode var. sclerophyllum (Hack.) Finot (7c) Trisetum caudulatum Trin. (8) Trisetum caudulatum Trin. var. caudulatum (8a) Trisetum caudulatum Trin. var. correae Nicora (8b) Trisetum cernuum Trin. (1) Trisetum dianthemum (Louis-Marie) Finot (9) Trisetum Jlavescens (L.) P. Beauv. (2) Trisetum foliosum Swallen (3) Trisetum irazuense (Kuntze) Hitchc. (4) Trisetum longiglume Hack. (10) Trisetum longiglume var. glabratum Nicora (10b) Trisetum longiglume Hack. var. longiglume (10a) Trisetum macbridei Hitchc. (11) Trisetum mattheii Finot (12) Trisetum nancaguense Finot (13) Trisetum oreophilum Louis-Marie (14) Volume 92, Number 4 2005 Trisetum Trisetum Trisetum Trisetum Trisetum Trisetum Trisetum oreophilum var. johnstonii Louis-Marie (14b) oreophilum Louis-Marie var. oreophilum (14a) phleoides (d'Urv.) Kunth (15) preslei (Kunth) E. Desv. (16) pyramidatum Louis-Marie ex Finot (17) spicatum (L.) K. Richt. (18) spicatum var. cumingii (Nees ex Steud.) Finot (18b) Trisetum spicatum (L.) K. Richt. var. spicatum (18a APPENDIX 2. INDEX TO SPECIMENS EXAMINED. NUMBER IN PARENTHESES CORRESPONDS TO NUMBER OF THE SPECIES IN THE TAXONOMIC TREATMENT AS INDICATED IN THE ALPHABETICAL INDEX (SEE APPENDIX 1). Alboff 1039 (15), s.n. 7 Mar. 1896 (1), s.n. Feb. 1896 (15); Ambrosetti & Mendez 407 (18b), 27457 (18b), 29874 (18b), 29875 (5), 29876 (18b), 29882 (18b), 29883 (5), 29884 (18b), 29885 (18b), 29889 (18b); Ameghino s.n. Dec. 1897 (18a); Anderson 387 (15); Andre 3907 (6); Anonymous 10 Feb. 1955 (16); Arancio 92129 (14b), 93021 (14b), s.n. 23 Mar. 1994 (14b); Arancio, Squeo & Leon 94043 (14b), 94250 (14b); Araya 16 (18a), 140 (8a); Arroyo 85159 (18b), 85160 (5), 85168 (18b); Arroyo et al. 215 (18a), 9276 (5), 92112 (5), 94454 (10a), 94477 (10a); Arroyo & Squeo 860055-a (18a); Arroyo, Boelcke, Gomez, Moore & Romanczuk 491 (15); Arroyo, Maldonado & Henriquez 91156 (16), 91158 (16); Arroyo, Mon & Romanczuk 200 (18b); Arroyo, Veloso & Pefialoza 870266 (18b); Arroyo, von Bohlen, Garcia & Gigoux 92112 (18b); Asplund 1028 (14a), 2010 (14a), 2057 (14a), 3585 (14a), 3798 (14a), 6023 (14a), 6156 (4), 6733 (4), 7215 (4), 7401 (6), 7441 (4), 7499 (6), 7547 (14a), 7896 (14a), 7978 (4), 8280 (4), 9686 (4), 11784 (11), 11793 (11), 16093 (4), 16131 (14a), 16868 (4), 17550 (14a). Baeza s.n. 9 Nov. 1913 (8a); Bailey s.n. Oct. 1958 (8a); Banks & Solander s.n. 1769 (15); Barrientos 224 (15); Barros 1987 (8a), 3982 (18a), 5706 (17), 5709 (17), 5719 (18a), s.n. 14 Nov. 1925 (8a), s.n. 24 Nov. 1925 (8a), s.n. 1 Nov. 1927 (8a); Bayer 4608 (13); Beck 13788 (14a); Beetle & Soriano HS-381a (10a); Behn s.n. 15 Dec. 1947 (17); Benoist 2389 (6), 4389 (6); Bertero 53 (8a), 996 (8a), 997 (8a), 998 (8a.); Bocher et al. 1894 (16), 1971 (16); Boelcke 1972 (16), 4132 (14b), 10764 (7a); Boelcke & Correa 5590 (18a), 5631 (18a), 5775 (18a), 5864 (8a), 6142 (8a), 6959 (7c), 6964 (7c); Boelcke & Hunziker 3657 (8b); Boelcke et al. 9778 (7c), 10011 (14b), 10039 (14b), 10061 (14b), 10927 (7a), 11033 (16), 11285 (16), 11318 (14b), 11452 1/2 (7a), 11674 (16), 13932 (10a), 14083 (7c); Boelcke, Bacigalupo, Correa 10226 1/2 (16); Boelcke, Correa, Bacigalupo et al. 10801 (18a); Bohlen & Cavieres 92312 (18b); Bonarelli 181 (17), 182 (17), 185 (17), s.n. 1 Mar. 1917 (5); Bridarolli 2147 (8a); Buchtien 6468 (14a), 8839 (14a), 8839 (18a), s.n. 14 Feb. 1903 (16), s.n. Feb. 1905 (17); Burkart 19905 (8a), 27451 (7a); Burkart & Tamayo 16733 (4); Burkart et al. 13930 (7c); Burmeister 3 (18a). Cabido s.n. 25 Jan. 1984 (18a); Cabrera 3662 (16), 5928 (18a), 11143 (18a), 11511 (1), 19754 (7c); Cabrera & Crisci 19145 (7c); Cabrera et al. 23024 (7c), 23170 (7c); Calderon & Rugolo 57 (7c), 72 (7c); Cafiulaf s.n. 29 Dec. 1946 (7a); Carrasco 266 (8a); Castellanos 1937 (15), s.n. 10 Feb. 1932 (1), s.n. 12 Jan. 1933 (18b), s.n. 28 Dec. 1933 (18a), s.n. 30 Jan. 1942 (5), s.n. 31 Jan. 1942 (18b), s.n. 31 Jan. 1942 (5), s.n. 6 Feb. 1942 (15); Castillo s.n. Feb. 1947 (16); Cazalet & Pennington 5740 (6); Correa 2953 (18b); Correa 4016 (18b); Correa et al. 5647 Finot et al. Trisetum in South America 565 (7c); Corte 34 (8a); Covas 101 (16); Cuatrecasas 9280 (18a), 20680 (4), 20847 (4); Cuming s.n. (18b); Cusato & Rossow 4524 (18b). Davalos 20 (4), 27 (18a); Davidse 3226 (3); Davies s.n. 17 Feb 1938 (15); Davis et al. 1381 (14a); Dawson 1285 (8a); Dawson & Schwabe 2481 (8a); Delfin s.n. Jan. 1887 (8a); Dessauer s.n. (8a); Diehl & Bravo 10843 (7a); Diem 458 (18b), 463 (2), 3229 (7c); Dimitri & Correa 176 (18b); Dimitri & Correa Luna 119 (18a); Dollenz 11 (18b), 1393 (18b); Dusen s.n. 5 Jan. 1897 (15), s.n. 19 Feb. 1905 (18a), s.n. Apr. 1937 (15); d'Urville 3 (15). Ehrenburg 49 (6), 137 (6); Eschscholtz s.n. (1); Eskuche & Klein 316 (8a), 1417-27 (8a), 1520-7 (7a); Eyerdam 10047 (18a), 10257 (18a); Eyerdam, Beetle & Grondona 24133 (18b). Fabris & Crisci 7608 (7a); Fabris & Solbrig 1171 (8b); Fagerlind & Wibon 833 (18a); Ferraro, Messuti & Vobis 4676 (15), 4703 (15); Finot & Baeza 1 (8a), 2 (8a), 7 (8a), 12 (18a), 14 (8a), 2069 (7a), 2070 (7a), 2071 (18a), 2073 (7a), 2074 (8a); Fonck 71 (8a); Fortunato 4823 & Elechosa (15). Gallinal, Aragone, Bergalli, Campal & Rosengurtt 5594 (2); Garaventa 2248 (8a), 4681 (7a), 6457 (8a); Gay 147 (8a), Gay 210 (18a), Gay s.n. (16), Gay s.n. (8a); Giaiotti et al. 23110 (16); Godoy 119 (1); Godoy, Hildebrand-Vogel & Vogel 3 (1); Gonzalez 766 (8b); Grondona 5695 (1), 7340 (15); Gunckel 630 (8a), 12426 (7a), 13797 (8a), 15101 (8a), 16027 (8a), 20295 (14b), 20297 (16), 20465a (14b), 21997 (8a), 40532 (8a), 44566 (8a), s.n. Feb. 1950 (16). Haenke s.n. (16); Harling 4537 (6); Hartweg 1449 (6); Hauman s.n. Feb. 1920 (7c); Hicken s.n. 30 Jan. 1912 (15), s.n. 30 Jan. 1912 (15); Hildebrand-Vogel 31 (1), 45 (1); Hitchcock 21059 (6), 22184 (14a), 22199 (14a), 22254 (14a), 22271 (14a), 22323 (14a), 22471 (14a), 22535 (14a); Holmgren 553 (6); Holm-Nielsen et al. 5553 (4); Hollermayer 26-a (9), 1252 (9), s.n. 14 Dec. 1927 (9); Hosseus 559 (7c), 560 (7c); Hunziker 8200 (15); Hutchinson 45 (8a), 59 (8a), 110 (8a), 212 (8a), s.n. 3 Jan. 1930, (10a). Illin 148 (7c), 169 (18a), s.n 1 Mar. 1900 (8a), s.n. 1901 (8a). Jaffuel 1806-a (18a), 1806-b (7a); James 189 (18a), 247 (18b); Jameson s.n. (6); Jiles 2952 (14b), 3649 (14b), 4136 (14b), 4246 (14b), 4694 (8a), 4801 (14b), 5888 (14b), 5889 (14b); Job 2431 (18a); Johnston 5407 (8a), 5539 (8a), 6097 (14b); Joseph 2948 (16); Junge s.n. 29 Oct. 1934 (8a). Kalela 1277 (8a), s.n. 27 Nov. 1937 (8a), s.n. 28 Dec. 1937 (8a); King s.n. 1826 (1); Koptaluti & Gomez 5816 (14b); Krapovickas 3933 (8a); Kurtz 7599 (16), 11086 (14b). Laegaard 101347 (14a), 101414 (4), 101604 (6); Laegaard et al. 103031 (4); Laegaard, Romoleroux & Leon 102731 (6); Lagiglia 2239 (16); Lagiglia & D'Antoni 1326 (16); Lahitte s.n. 7 Jan. 1964 (8b); Lahitte & Roquero 277 (8b), 363 (8a); Lahitte, Roquero & Lopez 67 (8a), 486 (8a); Lammers, Baeza & Penailillo 7894 (13); Landero 790 (5); Lara & Parker 41J (18a); Lechenby s.n. 31 May 1900 (2); Lechler 311 (8a), 1195 (18b), 1283 (1), 2846 (8a), 2948 (16); Leon & Calderon 1285 (8a), s.n. (7c), s.n. 17 Feb 1961 (8a); Lindig 1862 (4); Linnaeus s.n. (18a); Little & Paredes 68321E (6); Luti 1426 (15), 1640 (15). Macbride & Featherstone 1131 (11); Magens 3032 (18b), 3197 (18b), 3428 (18b), 3433 (18b); Mandon 1309 (14a), 1857 (14a); Marin 1432 (14a); Marticorena & Matthei 664 (16); Marticorena et al. 83462-B (14b); Martico- 566 Annals of the Missouri Botanical Garden rena, Matthei & Quezada 86 (12); Marticorena, Weldt & Crisci 1967 (9), 1982 (8a); Martinez Carretero 1272 (16); Matthei & Bustos 111 (1); Matthei 174 (8a); Mendez & Ambrosetti s.n. 5 Feb. 1978 (18b); Mendez & Willoud c-320-7196 (16); Molau & Eriksen 3297 (6); Montero 538 (8a), 1354 (8a), 1962 (8a), 4487 (7a), 4497 (7a), 4508 (8a), 4956 (8a), 9557 (18a); Mooney, H. & B. Mooney 546 (16); Moore 848 (15); Moore & Goodall 398 (18b); Mufioz 2709 (8a), 2717 (8a), 3981 (14b), 4104 (18b), 4144 (18b), s.n. (18a); Munoz, M. et al. 3491 (16); Munoz & Coronel 1232 (8a); Mufioz & Johnson 2524 (8a), 2583 (8a), 2630 (8a), 2635 (8a), 2676 (8a), 2732 (8a), 2737 (8a), 3211 (8a); Mufioz & Schick 1493 (8a), 1545 (8a), 2558 (8a), 2562 (8a). Neger s.n. Nov. 1896 (8a); Nicora 7432 (7c), 7478 (8a), 9344 (18a), 9485 (18a), 9519 (18a), 9603 (18a), 9610 (9), 10081 (18a), 10235 (18a), 10284 (18a), 10344 (18a), 10348 (18a), Nowak & Marcillo 79 (6). Offermann s.n. 18 Jan. 1930 (8a); Ortega s.n. Feb. 1879 (15). Paci 15731 (7c); Panza s.n. 25 Dec. 1970 (18a); Parodi 2700 (7c), 3196 (16), 10122 (14a), 10127 (14a), 11827 (1), 15321 (7c), 15560 (18b), 15582 (18b), 15611 (8a); Pedersen 1468 (8a), 1523 (1), s.n. 6 Feb. 1952 (8a); Penland 584 & Summers (14a); Pennington 450b (15); Pefialoza, Claros, Cavieres & Flores 91121 (16), 91122 (16), 91126 (16); Perez Moreau 144 (14b, pro parte), 1949 (1), 30-258 (14b), s.n. 29 Jan. 1941 (8b), s.n. 21 Jan. 1942 (18b), s.n. 14 Jan. 1949 (8a), s.n. 1949 (7c); Perez Moreau & Guarreva s.n. 2 Feb. 1948 (1); Perez-Moreau & Perrone s.n. 12 Feb. 1950 (14b); Perrone s.n. 26 Feb. 1950 (14b); Peterson & Refulio-Rodrfguez 15135 (4); Peterson et al. 9146 (4); Peterson, Annable & Poston 8765 (18a), 8866 (14a); Peterson, Judziewicz & King 9075 (6); Peterson, Judziewicz, King & j0rgensen 9176 (14a); 9239 (14a); Peterson, Soreng & Laegaard 13173 (18a); Pettersson 15 (18a), 41 (14a), 57 (14a); Pfister 370 (8a); s.n. (18b), s.n. 20 Oct.1943 (8a), s.n. 12 Nov. 1944 (8a), s.n. 26 Nov. 1944 (8a), s.n. 17 Jan. 1945 (7b), s.n. 10 Dec. 1946 (8a), s.n. 5 Jan. 1947 (8a), s.n. 5 Jan. 1947 (8b), s.n. 10 Jan. 1947 (7a), s.n. 12 Jan. 1947 (7a), s.n. 1 Jan. 1948 (7a), s.n. 9 Jan. 1949 (7a), s.n. 10 Oct. 1951 (8a), s.n. 14 Jan. 1953 (18a); Pfister & Ricardi s.n. 31 Dec. 1951 (15), s.n. 10 Jan. 1952 (18a); Philippi 218 (7a), 229 (7a), s.n. SGOPHIL 220 (8a), s.n. SGO-PHIL-235 (15), s.n. Jan. 1855 (8a), s.n. Feb. 1858 (8a), s.n. Jan. 1860 (8a), s.n. Nov. 1861 (8a), s.n. 1864/65 (18b), s.n. Jan. 1877 (8b), s.n. SGO (8a), s.n. SGO (8b); Philippi, F. s.n. (18b), s.n. Oct. 1872 (8a), s.n. Jan. 1877 (7a), s.n. Jan. 1877 (16), s.n. Jan. 1878 (16); Philippi, O. s.n. Jan. 1887 (9); Pico s.n. 9 Jan. 1919 (15); Pisano 2451 (15); Pisano, Henriquez & Dominguez 6941 (15), 6962 (15), 7396 (15), 7436 (18b), 7463 (18b), 7543 (15), 7566 (15); Pittier 1435 (4); Poeppig s.n. 1828 (7), s.n. BAA 3364 (8a). Ragonese 234 (7a); Reiche s.n. (7b), s.n. Mar. 1899 (16); Remy s.n., Oct. 1856 (6); Ricardi 5604 (7a), s.n. 11 Dec. 1950 (8a); Ricardi & Marticorena 5723/1884 (7a), 5812/1973 (7a); Ricardi & Matthei 3 (7a), 15 (7a), 27 (8a), 166 (8a), 177 (18a), 184 (18b), 236 (8a), 263 (18b), 316 (15), 335 (1), 385 (18b); Ricardi, Marticorena & Matthei 1829 (8a), 1976 (8a); Ricardi, Marticorena & Torres s.n. 2 Nov. 1957 (8a); Riggi 27/1052 (18b); Robinson & Beltran 3041 (4); Roig 1 (7c), 50 (14b), 11950 (14b), 12356 (6), 15589 (18b); Roig et al. 103 (5); Roig et al. 444 (15); Roig J. 12000 (14a), s.n. 5 Feb. 1986 (15); Roig Jufient 13353 (18b), Roig Jufient, E, C. Roig Jufient & F. A. Roig 14912 (18b); Roig, A. & F. Roig 14980 (18b); Roig, An- chorena, Mendez & Ambrosetti 115 (18b); Roig, F, C. Roig & F. A. Roig 14911 (15); Roig, Roig Jufient & Martinez Carretero 14266 (18a); 14748 (18b); Roquero 31 (18a), 132 (18b), 169 (18a), 297 (18a), 326 (8a), 434 (18a), 452 (8a); Rue s.n. Nov 1958 (15); Rugolo 1188 (8a). 1283 (8a); Rugolo & Agrasar 315 (7c), 318 (7c), 570 (7c); Rugolo de Agrasar 233 (8b), 451 (7a), 1093 (18a), 1176 (18a), 1234-1 (7a), 1234-2 (7c), 1252 (18b); Rugolo de Agrasar & Agrasar 148 (7a), 573 (10b), 5846 (16); Rugolo de Agrasar et al. 12381 (1); Ruiz Leal 79 (16), 3616 (7c), 6625 (16), 16886 (7c); Ruiz Leal & Roig 15023 (1); Rumboll 120 (18b). Saavedra & Pauchard 6 (16); Sagastegui, Mostacero & Leiva 12056 (14a); Salaverry 17 (14a); Sanchez-Vega et al. 1380 (14a); Sanchez-Vega, Torrel, & Medina 2557 (14a), Sanchez-Vega, Molau & Ohman 3800 (14a), 3812 (14a); Sanchez-Vega & Castillo 6370 (14a); Sanchez-Vega & Cabanillas 6790 (14a); Schajowski 55-a (18a), 86 (7c), 133 (7c), 134 (7c), 178 (7c); Schlegel 2371 (8a), 3475 (7a), 3537 (7a), 3645 (8a), 4065 (8a), 7187 (1), 7194 (1), 7322 (18a), 7523 (18a), 8070 (1), 8122 (1), s.n. Mar. 1958 (8a); Schwabe 69 (18a); Schwing s.n. 23 Jun. 1932 (2); Sklenar & Kosteckova 80-12 (6), 812 (6); Skottsberg 69 (15), 236 (15), s.n. Mar 1902 (15); Smith, Valencia & Gonzales 9772 (11); Smith, Valencia & Minaya 9943 (14a); Smith, Valencia, Gonzalez & Buddensiek 12224 (14a); Sneidern 3027 (4); Sodiro 1893 (14a), Sodiro s.n. (18a), s.n. Oct. 1891 (4); Solomon 13221 (18a); Soriano 2409 (8a), 2490 (8a), 3184 (18a), 4334 (8b), 4352 (7c), 4847 (5), s.n. Feb. 1943 (18a); Sparre 1678 (16), 4873 (7a); Sparre & Constance 10778 (18a); Spegazzini s.n. (18a), s.n. Mar. 1882 (15), s.n. Dec. 1908 (16); Stebbins 9061 (13); Steyermark 57482 (3), 62083 (4), 62488 (4); Swallen 7064 (14a). T.B.P.A. 2404 (18b), 3117 (18a), 3119 (5), 3140 (18a); Teillier 1536 (14b); Teillier et al. 2548 (16); Tovar 2474 (14a), 2539 (14a), 2933 (14a); Trombotto s.n. 19-22 Feb. 1984 (14b); Trombotto & Ahumada 11105 (14b); Tsujii 172 (15). Valla et al. 3065 (18a); Vargas 7045 (14a); Vellerini 257 (8a), 265 (8a), 297 (8a); Vidal 233 (18a), 234 (18a); Vidal Gormaz 265 (10a), s.n. SGO 37066 (8a), s.n. SGO-PHIL239b (9); Villagran & Leiva 7525 (18b); Villagran & Mesa 414 (8a); Villagran et al. 8462 (16); Villagran, Aguila & Leiva 6954 (8a); Villamil 6 (7a), 8396 (15); Villamil et al. 2844 (18b); Villarroel & Weldt 151 (13); Virlet 1382 (4); Vogel 5 (1), 519 (1), 540 (1), 545 (18a), s.n. (1). Werdermann 655 (18a). Zoellner 5859 (16), 13135 (8a), 18030 (8a), 18196 (8b), s.n. 2 Feb. 1968 (7a), s.n. 14 Feb. 1992 (7a). APPENDIX 3. LECTOTYPIFICATIONS OF SUBGENERIC TAXA. Trisetum sect. Anaulacoa Louis-Marie, Rhodora 30(359): 211, 212. 1928. TYPE: Trisetum flavescens (L.) P. Beauv., lectotype, designated here. Trisetum sect. Aulacoa Louis-Marie, Rhodora 30(359): 211. 1928; 30(360): 243. 1929. TYPE: Trisetum floribundum Pilg., lectotype, designated here. Trisetum subg. Heterolytrum Louis-Marie, Rhodora 30(359): 211, 212. 1928. TYPE: Trisetum flavescens (L.) P. Beauv., lectotype, designated here. Trisetum subg. Isolytrum Louis-Marie, Rhodora 30(259): 211. 1928; 30(360): 244. 1929. TYPE: Trisetum longiglume Hack., lectotype, designated here. Trisetum subsect. Koeleriformia Louis-Marie, Rhodora 30(359): 211. 1928; 30(360): 241. 1929. TYPE: Volume 92, Number 4 2005 Trisetum micratherum E. Desv., lectotype, designated here. Trisetum subsect. Spheiiopholidea Louis-Marie, Rhodora 30(259): 211. 1928; 30(360): 240. 1929. TYPE: Trisetum interruptum Buckl., lectotype, designated here. APPENDIX 4. LIST OF NAMES AND SYNONYMS OF ALL TAXA MENTIONED IN THIS MANUSCRIPT. Accepted names are presented in bold and synonyms are italicized. Acrospelion Besser ex Schult. & Schult. f. = Trisetum Aira spicata L. = Trisetum spicatum var. spicatum Aira subspicata L. = Trisetum spicatum var. spicatum Amphibromus Nees Arrhenatherum P. Beauv. Avena L. Avena airoides Koel. = Trisetum spicatum var. spicatum Avena cernua (Trin.) Kunth = Trisetum cernuum Avena flavescens L. = Trisetum flavescens Avena leptostachys Hook. f. = Trisetum cernuum Avena phleoides d'Urv. = Trisetum phleoides Avena pilosa J. Presl = Trisetum preslei Avena preslei Kunth = Trisetum preslei Bromus L. Bromus berteroanus Colla Bromus trinii E. Desv. = Bromus berteroanus Calamagrostis irazuensis Kuntze = Trisetum irazuense Deschampsia P. Beauv. Deschampsia airiformis (Steud.) Benth. & Hook.f. Deschampsia andicola (Louis-Marie) Valencia = Trisetum longiglume var. longiglume Deschampsia lasiantha Phil. = Trisetum preslei Dielsiochloa Pilg. Dielsiochloa floribunda (Pilg.) PilgGraphephorum Desv. Helictotrichon Besser ex Schult. & Schult. f. Helictotrichon virescens (Nees ex Steud.) Henrard Koeleria Pers. Koeleria caudulata (Trin.) Griseb. = Trisetum caudulatum var. caudulatum Koeleria cumingii Nees ex Steud. = Trisetum spicatum var. cumingii Koeleria fueguina C. E. Calderon ex Nicora Koeleria lechleri Steud. = Trisetum caudulatum var. caudulatum Koeleria spicata Reichb. ex Willk. & Lange = Trisetum spicatum var. spicatum Koeleria subspicata (L.) Rchb. = Trisetum spicatum var. spicatum Leptophyllochloa C. E. Calderon Leptophyllochloa micrathera (E. Desv.) C. E. Calderon Peyritschia E. Fourn. Peyritschia conferta (Pilg.) Finot Peyritschia deyeuxioides (Kunth) Finot Raimundochloa A. M. Molina Rebentischia Opiz = Trisetum Rebentischia flavescens Opiz = Trisetum flavescens Relchela Steud. Rostraria Trin. Rupestrina Prov. = Trisetum Finot et al. Trisetum in South America 567 Rupestrina pubescens Prov. = Trisetum spicatum var. spicatum Sphenopholis Scribn. Trisetaria airoides (Koeler) Baumg. = Trisetum spicatum var. spicatum Trisetaria flavescens (L.) Baumg. = Trisetum flavescens Trisetum Pers. Trisetum subg. Deschampsioidea (Louis-Marie) Finot Trisetum subg. Heterolytrum Louis-Marie = Trisetum subg. Trisetum Trisetum subg. Isolytrum Louis-Marie = Trisetum sect. Trisetaera Trisetum subg. Trisetum Trisetum sect. Anaulacoa Louis-Marie = Trisetum sect. Trisetum Trisetum sect. Aulacoa Louis-Marie = Dielsiochloa Trisetum sect. Carpatica Chrtek Trisetum sect. Trisetaera Asch. & Graebn. Trisetum sect. Trisetum Trisetum subsect. Carpatica (Chrtek) Prob. = Trisetum sect. Carpatica Trisetum subsect. Deschampsioidea Louis-Marie = Trisetum subg. Deschampsioidea Trisetum subsect. Graphephorum (Desv.) Louis-Marie = Graphephorum Trisetum subsect. Koeleriformia Louis-Marie = Leptophyllochloa Trisetum subsect. Spenopholidea Louis-Marie = Sphenopholis Trisetum airiforme Steud. = Deschampsia airiformis Trisetum airoides (Koeler) P. Beauv. ex Roem. & Schult. = Trisetum spicatum var. spicatum Trisetum albidum Sodiro = Trisetum spicatum var. spicatum Trisetum ambiguum Rugolo & Nicora Trisetum andicola Louis-Marie = Trisetum longiglume var. longiglume Trisetum andinum Benth. Trisetum araeanthum Phil. = Leptophyllochloa micrathera Trisetum barbatum Steud. = Bromus berteroanus Colla Trisetum barbinode Trin. Trisetum barbinode Trin. var. barbinode Trisetum barbinode var. hirtiflorum (Hack.) LouisMarie Trisetum barbinode var. sclerophyllum (Hack, ex Stuck.) Finot Trisetum biflorum Phil. = Trisetum dianthemum Trisetum brachyatherum Phil. = Leptophyllochloa micrathera Trisetum [?] brasiliense Louis-Marie = Deschampsia brasiliensis (Louis-Marie) Valencia Trisetum buchtienii Hack. = Trisetum preslei Trisetum caudulatum Trin. Trisetum caudulatum Trin. var. caudulatum Trisetum caudulatum var. correae Nicora Trisetum cernuum Trin. Trisetum chiloense Phil. = Trisetum caudulatum var. caudulatum Trisetum chromostachyum E. Desv. = Trisetum caudulatum var. caudulatum Trisetum confertum Pilg. = Peyritschia conferta Trisetum cumingii (Nees ex Steud.) Nicora = Trisetum spicatum var. cumingii 568 Trisetum cumingii var. cumingii = Trlsetum spicatiun var. cumingii Trisetum cumingii var. santacrucense Nicora = Trisetum spicatum var. cumingii Trisetum depauperatum Phil. = Leptophyllochloa micrathera Trisetum deyeuxioides (Kunth) Kunth Trisetum dianthemum (Louis-Marie) Finot Trisetum erectum Phil.•name of uncertain application Trisetum evolutum (E. Fourn.) Hitchc. = Peyritschia deyeuxioides Trisetum flavescens (L.) P. Beauv. Trisetum flavescens subsp. pratense (Pers.) Asch. & Graebn. = Trisetum flavescens Trisetum floribundum Pilg. = Dielsiochloa floribunda Trisetum foliosum Swallen Trisetum fournieranum Hitchc. = Trisetum irazuense Trisetum fraudulentum Steud. = Trisetum cernuum Trisetum heterogamum Steud. ex Lechler = Trisetum caudulatum var. caudulatum Trisetum heteronymum Steud. = Trisetum caudulatum var. caudulatum Trisetum hirsutum Phil. = Trisetum phleoides Trisetum hirtiflorum Hack. = Trisetum barbinode var. hirtiflorum Trisetum hirtum Trin. = Bromus berteroanus Colla Trisetum irazuense (Kuntze) Hitchc. Trisetum [?] juergensii Hack. Trisetum lasiolepis E. Desv. = Trisetum preslei Trisetum laxiflorum Phil. = Leptophyllochloa micrathera Trisetum laxum Phil. = Leptophyllochloa micrathera Trisetum lechleri (Steud.) Nicora = Trisetum caudulatum var. caudulatum Trisetum longiglume Hack. Trisetum longiglume var. glabratum Nicora Trisetum longiglume Hack. var. longiglume Trisetum macbridei Hitchc. Trisetum malacophyllum Phil. = Trisetum spicatum var. cumingii Trisetum malacophyllum Steud. = Trisetum caudulatum var. caudulatum Trisetum mattheii Finot Trisetum micratherum E. Desv. = Leptophyllochloa micrathera Trisetum mollifolium Louis-Marie = Trisetum spicatum var. cumingii Trisetum monticola Phil. = Trisetum caudulatum var. caudulatum Trisetum nancaguense Finot Trisetum nemorosum Phil. = Leptophyllochloa micrathera Trisetum ochrostachyum Phil. = Trisetum caudulatum var. caudulatum Trisetum oreophilum Louis-Marie Trisetum oreophilum var. johnstoni Louis-Marie Trisetum oreophilum var. oreophilum Trisetum paradoxum Phil.•name of uncertain application Trisetum phleoides (d'Urv.) Kunth Trisetum pratense Pers. = Trisetum flavescens View publication stats Annals of the Missouri Botanical Garden Trisetum preslei (Kunth) E. Desv. Trisetum preslei var. buchtienii (Hack.) Louis-Marie = Trisetum preslei Trisetum preslei var. lasianthum (Phil.) Louis-Marie = Trisetum preslei Trisetum pyramidatum Louis-Marie ex Finot Trisetum rosei Scribn. & Merr. Trisetum scabriflorum Hitchc. = Trisetum irazuense Trisetum sclerophyilum Hack. = Trisetum barbinode var. sclerophyilum Trisetum pyramidatum Louis-Marie ex Finot Trisetum spicatum (L.) K. Richt. Trisetum spicatum var. cumingii (Nees ex Steud.) Finot. Trisetum spicatum K. Richt. var. spicatum Trisetum spicatum subsp. andinum (Renth.) Hulten = Trisetum andinum Trisetum spicatum subsp. bolivianum Hulten = Trisetum spicatum var. spicatum Trisetum spicatum var. phleoides (d'Urv.) Hack. = Trisetum phleoides Trisetum spicatum subsp. phleoides (d'Urv.) Hulten = Trisetum phleoides Trisetum spicatum subsp. phleoides (d'Urv.) Macloskie = Trisetum phleoides Trisetum spicatum var. andinum (Renth.) Louis-Marie = Trisetum andinum Trisetum spicatum var. dianthemum Louis-Marie = Trisetum dianthemum Trisetum spicatum var. hirsutum Louis-Marie = Trisetum phleoides Trisetum splendidulum Steud. = Trisetum caudulatum var. caudulatum Trisetum subspicatum (L.) P. Reauv. = Trisetum spicatum var. spicatum Trisetum subspicatum subsp. phleoides (d'Urv.) Hack. = Trisetum phleoides Trisetum tomentosum (E. Desv.) Nicora = Koeleria fueguina Trisetum trinii (E. Desv.) Louis-Marie = Bromus berteroanus Colla Trisetum variabile E. Desv. = Trisetum caudulatum var. caudulatum Trisetum variabile E. Desv. var. variabile = Trisetum caudulatum var. caudulatum Trisetum variabile subsp. virescens Macloskie = (discussed under T. caudulatum var. caudulatum) Trisetum variabile var. chiloense (Phil.) Louis-Marie = Trisetum caudulatum var. caudulatum Trisetum variabile vox. flavescens E. Desv. = Trisetum caudulatum var. caudulatum Trisetum variabile var. intonsum Louis-Marie = Helictotrichon virescens (Nees ex Steud.) Henrard Trisetum variabile var. vidalii (Phil.) Louis-Marie = Trisetum caudulatum var. caudulatum Trisetum variabile var. virescens E. Desv. = Trisetum caudulatum var. caudulatum Trisetum vidalii Phil. = Trisetum caudulatum var. caudulatum Trisetum virescens Nees ex Steud. = Helictotrichon virescens (Nees ex Steud.) Henrard Trisetum weberbaueri Pilg. = Dielsiochloa floribunda

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