A Revision of Deschampsia, Avenella, and Vahlodea (Poaceae, Poeae, Airinae) in South America1

Jorge Chiapella; Fernando O. Zuloaga

A revision of the species of Deschampsia P. Beauv., Avenella (Bluff & Fingerh.) Drejer, and Vahlodea Fr. (Poaceae, Poeae, Airinae) present in South America is given. Fifteen species of Deschampsia and two monospecific genera segregated from Deschampsia—Avenella and Vahlodea—are found in the Andes of Argentina and Chile south of 30uS, typically in humid or damp sites and wetlands. Isolated populations of the cosmopolitan D. cespitosa (L.) P. Beauv. are also found in Bolivia and highlands of southern Brazil. Keys to differentiate among the species of Deschampsia and the allied genera Avenella and Vahlodea are provided. A lectotype is designated for D. berteroana (Kunth) Trin. RESUMEN Se presenta una revisión de las especies de Deschampsia P. Beauv., Avenella (Bluff & Fingerh.) Drejer y Vahlodea Fr. de Sudamérica. Se hallaron quince especies de Deschampsia y los dos géneros monotípicos Avenella y Vahlodea (segregados de Deschampsia), típicamente en lugares hú medos o pantanosos de los Andes de Argentina y Chile al sur de los 30uS. Poblaciones aisladas de la especie cosmopolita D. cespitosa (L.) P. Beauv. se encuentran en Bolivia y las tierras altas del sur de Brasil. Se dan claves para diferenciar Deschampsia de sus géneros aliados Avenella y Vahlodea y de las especies de Deschampsia. Se designa el lectotipo de D. berteroana (Kunth) Trin. Deschampsia P. Beauv. consists of about 30 species found in cold temperate regions of both hemispheres

Volume 97 Number 2 2010 Annals of the Missouri Botanical Garden A REVISION OF DESCHAMPSIA, AVENELLA, AND VAHLODEA (POACEAE, POEAE, AIRINAE) IN SOUTH AMERICA1 Jorge Chiapella2 and Fernando O. Zuloaga3 ABSTRACT A revision of the species of Deschampsia P. Beauv., Avenella (Bluff & Fingerh.) Drejer, and Vahlodea Fr. (Poaceae, Poeae, Airinae) present in South America is given. Fifteen species of Deschampsia and two monospecific genera segregated from Deschampsia—Avenella and Vahlodea—are found in the Andes of Argentina and Chile south of 30uS, typically in humid or damp sites and wetlands. Isolated populations of the cosmopolitan D. cespitosa (L.) P. Beauv. are also found in Bolivia and highlands of southern Brazil. Keys to differentiate among the species of Deschampsia and the allied genera Avenella and Vahlodea are provided. A lectotype is designated for D. berteroana (Kunth) Trin. RESUMEN Se presenta una revisión de las especies de Deschampsia P. Beauv., Avenella (Bluff & Fingerh.) Drejer y Vahlodea Fr. de Sudamérica. Se hallaron quince especies de Deschampsia y los dos géneros monotı́picos Avenella y Vahlodea (segregados de Deschampsia), tı́picamente en lugares húmedos o pantanosos de los Andes de Argentina y Chile al sur de los 30uS. Poblaciones aisladas de la especie cosmopolita D. cespitosa (L.) P. Beauv. se encuentran en Bolivia y las tierras altas del sur de Brasil. Se dan claves para diferenciar Deschampsia de sus géneros aliados Avenella y Vahlodea y de las especies de Deschampsia. Se designa el lectotipo de D. berteroana (Kunth) Trin. Key words: Airinae, Aveneae, Avenella, Deschampsia, Poaceae, Poeae, Vahlodea. Deschampsia P. Beauv. consists of about 30 species found in cold temperate regions of both hemispheres (Hultén, 1941; Parodi, 1949; Hitchcock et al., 1969; Bor, 1970; Tzvelev, 1976; Cronquist et al., 1977; Clarke, 1980; Clayton & Renvoize, 1986; Beetle, 1987; Conert, 1987; Rzedowski & Rzedowski, 1990; Hickman, 1993). The number of species may increase if several subspecies of D. cespitosa (L.) P. Beauv. 1 This paper is part of the doctoral thesis of J.C. at the Universidad Nacional del Comahue, Bariloche, Argentina. Curators of B, BA, BAA, BAB, BM, CONC, CORD, K, LE, P, SGO, S, SI, US, and W helped in finding type specimens and representative specimens. C. Ezcurra provided constant support throughout the realization of this work. J.C. acknowledges fellowships from the Universidad Nacional del Comahue (Argentina), the Österreichischer Austauschdienst (ÖAD-Austria), and a Kew Latin American Research Fellowship (KLARF), Royal Botanic Gardens, Kew, which allowed visits to B, BM, K, LE, P, S, and W. Comments by P. M. Peterson and V. Hollowell improved the manuscript. 2 Laboratorio Molecular, IMBIV, Universidad Nacional de Córdoba, Vélez Sarsfield 1609, X5016GCA Córdoba, Argentina. jchiapella@imbiv.unc.edu.ar 3 Instituto de Botánica Darwinion, Labardén 200, B1642HYD San Isidro, Argentina. doi: 10.3417/2008115 ANN. MISSOURI BOT. GARD. 97: 141–162. PUBLISHED ON 00 MONTH 2010. Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:46 141 Cust # 2008115 142 Annals of the Missouri Botanical Garden from Asia and the North Pacific are raised to specific status (Chiapella & Probatova, 2003). Some endemics exist in high mountains or islands, most notably D. chapmanii Petrie, D. gracillima Kirk, D. pusilla Petrie, and D. tenella Petrie in New Zealand; D. christophersenii C. E. Hubb., D. mejlandii C. E. Hubb., D. robusta C. E. Hubb., and D. wacei C. E. Hubb. in the Tristan da Cunha Islands; D. klossii Ridl. in the mountains of Indonesia and Papua New Guinea; D. argentea (Lowe) Lowe and D. maderensis (Hack. & Bornm.) Buschm. in the Madeira Islands; D. domingensis Hitchc. & Ekman in the Cordillera Central of Hispaniola; D. foliosa Hack. in the Azores Islands; D. koelerioides Regel and D. pamirica Roshev. in the Pamir Mountains; D. liebmanniana (E. Fourn.) Hitchc. in the mountains of Central Mexico; D. nubigena Hillebr. in Hawaii; and D. angusta Stapf & C. E. Hubb. in the high mountains of tropical Africa. The species present in South America are restricted to the subcontinent, with the exception of D. danthonioides (Trin.) Munro ex Benth. and D. elongata (Hook.) Munro ex Benth., which are also found in western North America; the nearly cosmopolitan D. cespitosa; and D. setacea (Huds.) Hack., which is found in Europe and central Chile. The genus Deschampsia was described by Palisot de Beauvois (1812) to honor J. C. A. Loiseleur Deslong Champs, physician of the rescue expedition of the explorer La Pérouse. Palisot based his new genus on Aira cespitosa L., a species described in the first edition of Species Plantarum (Linnaeus, 1753), and included three other taxa (D. discolor Roem. & Schult., D. juncea P. Beauv., and D. parviflora P. Beauv.). Linnaeus (1753: 64) separated the species of Aira into two groups, one with muticous lemmas and the other with awned lemmas, and included A. cespitosa in the latter group. In addition to this character, Palisot (1812: 91) characterized Deschampsia as having paniculate inflorescences; 2- or 3-flowered ‘‘glumes’’ (5 spikelets); glumes longer than the spikelets; lemmas with several teeth; and awn straight, inserted in or near the base of the lemmas, and slightly longer than the lemmas. This rather vague combination of characters and the high morphological diversity and abundance of Deschampsia, particularly of D. cespitosa in Eurasia (Chiapella, 2000; Chiapella & Probatova, 2003), resulted in a weak generic concept, as the same description applies to several slightly different forms. Avenella flexuosa (L.) Drejer and Vahlodea atropurpurea (Wahlenb.) Fr. ex Hartm. have had an obscure taxonomic history, being neglected in nearly all European, North American, and South American revisions (see Albers, 1972a, b, 1978, 1980a, b, c; Albers & Butzin, 1977; Garcı́a-Suárez et al., 1997; Frey, 1999), and instead treated under Deschampsia as D. atropurpurea (Whalenb.) Scheele and D. flexuosa (L.) Trin. A molecular sequence study (Chiapella, 2007) has shown that Deschampsia is monophyletic, and, despite a close relationship with the main core of Deschampsia (where the South American species are included), neither Avenella (Bluff & Fingerh.) Drejer or Vahlodea Fr. are included and should be considered as separate genera. The first South American descriptions of taxa later treated in Deschampsia and Vahlodea were done by Hooker (1847), who described Aira antarctica Hook. f., A. kingii Hook. f., A. magellanica Hook. f., and A. parvula Hook. f. Hooker also mentioned collections from southern Patagonia (Port Famine, Strait of Magellan) and the Falkland Islands (Islas Malvinas) of another common European grass, Aira flexuosa L. (; Avenella flexuosa (L.) Drejer), asserting that these plants could not be distinguished from the European plants. No molecular study has addressed whether the South American populations of Avenella flexuosa, Deschampsia cespitosa, and Vahlodea atropurpurea are introduced or native to the region (collection dates in additional specimens studied indicate localities where collecting took place before significant human influence could be considered the likely cause of introduction). Desvaux (1854) made three new combinations: Deschampsia antarctica (Hook. f.) E. Desv., D. kingii (Hook. f.) E. Desv., and D. parvula (Hook. f.) E. Desv. Other species mentioned by Desvaux (1854) for southern Patagonia and the Andes of central Chile were D. flexuosa (; Avenella flexuosa (L.) Drejer), D. discolor (5 D. setacea (Huds.) Hack.), and D. pulchra Nees & Meyen (; D. cespitosa var. pulchra (Nees & Meyen) Nicora); another species, Aira magellanica, was reduced to the synonymy of D. atropurpurea (Whalenb.) Scheele (; Vahlodea atropurpurea (Wahlenb.) Fr. ex Hartm.). The new genus Monandraira E. Desv., also described by Desvaux (1854) on the basis of plants with single staminate flowers, comprised two taxa from the Andes of central Chile: Monandraira berteroana (Kunth) E. Desv. (; D. berteroana (Kunth) Trin.) and M. glauca E. Desv. (5 D. danthonioides (Trin.) Munro ex Benth.). The prolific German-born Chilean botanist, R. A. Philippi, described Deschampsia latifolia Phil. (non D. latifolia Hoechst. ex A. Rich.) and D. laxa Phil. in 1858; D. lasiantha Phil. (5 Trisetum preslei (Kunth) E. Desv.) in 1864; D. andina Phil. (5 D. cespitosa (L.) P. Beauv.) and Monandraira patula Phil. (; D. patula (Phil.) Pilg. ex Skottsb.) in 1873; and D. brachyphylla Phil. (5 Vahlodea atropurpurea (Wahlenb.) Fr. ex Hartm.), D. fuegina Phil. (5 D. antarctica E. Desv.), D. martinii Phil. (5 Avenella flexuosa (L.) Drejer), and D. micrantha Phil. (see Excluded or Uncertain Taxa) in 1896. The beginning of the century brought the first Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:47 142 Cust # 2008115 Volume 97, Number 2 2010 Chiapella & Zuloaga Revision of Deschampsia, Avenella, and Vahlodea assessments of the genus, as the expeditions to Patagonia by Dusén (1900) and Macloskie (1904) produced the first general assessments of Deschampsia in the region, recognizing five and 12 species, respectively. Hauman (1918) described D. cordillerarum Hauman, pointing out its resemblance to D. flexuosa (; A. flexuosa) but noting that it differs by xerophilous characters (dense tufts of short, rigid leaves), longer ligules, and smaller spikelets. Valencia (1941) transferred four species of Trisetum Pers. to Deschampsia and made the new combinations D. juergensii (Hack.) Valencia (; T. juergensii Hack.), D. conferta (Pilg.) Valencia (; Peyritschia conferta (Pilg.) Finot), D. brasiliensis (Louis-Marie) Valencia (see Excluded or Uncertain Taxa), and D. andicola (Louis-Marie) Valencia (5 T. longiglume Hack.). Parodi (1949) rejected these combinations, arguing that the structure of the lemmas with awns derived from the middle nerve, as displayed by these taxa, corresponds more to Trisetum; Parodi also updated Deschampsia in South America, accepting 17 species, including three new species and two varieties (D. looseriana Parodi, D. looseriana var. triandra Parodi, D. mendocina Parodi, D. venustula Parodi, D. berteroana var. parvispicula Parodi) and three new combinations (D. glauca (E. Desv.) Parodi, D. elegantula (Steud.) Parodi, D. elegantula var. patula (Phil.) Parodi). 2a. Leaf blades filiform, conduplicate, convolute, folded or bristlelike, ca. 1 mm wide; lemma awns slender . . . . . . . . . . . . . . . . . . I. Avenella 2b. Leaf blades flat, never conduplicate or convolute, up to 6 mm wide; lemma awns stout . . . . . . . . . . . . . . . . . . . . . . II. Vahlodea 1b. Spikelets lanceolate, 2–4 mm; ligules acute, 5–10(12) mm; plants annual or perennial . . . . . . III. Deschampsia MATERIALS AND METHODS This study is based on herbarium specimens of B, BA, BAA, BAB, BM, CONC, CORD, K, LE, P, S, SGO, SI, US, and W (Holmgren et al., 1990), including nearly all types. In our study, we used the phylogenetic species concept (Cracraft, 1983), whose main assumptions are: a species is the smallest diagnosable unit of (sexual) populations possessing at least one diagnostic character that is present in all members of the group (Cracraft, 1983; Nixon & Wheeler, 1990; Quicke, 1993). In the present work, both qualitative and quantitative characters were considered diagnostic, although it was not always possible to find clear gaps in quantitative characters. In the descriptions of leaves, the first measurement range is the blade length (expressed in centimeters) and the second measurement range is the blade width (expressed in millimeters). KEY FOR DISTINGUISHING AVENELLA, DESCHAMPSIA, SOUTH AMERICA AND VAHLODEA IN 1a. Spikelets ovoid to oblong, 3.5–5 mm; ligules ovate to obtuse, blunt, never acute, 2.5–5 mm; plants always perennial. 143 TAXONOMIC TREATMENT I. Avenella (Bluff & Fingerh.) Drejer, Fl. Excurs. Hafn. 32. 1837. TYPE: Avenella flexuosa (L.) Drejer. Erioblastus Honda, Rep. Fl. Mt. Daisetsu 12: 73. 1930. TYPE: Erioblastus flexuosus Honda ex Nakai. Plants perennial, caespitose, sometimes with short rhizomes, flowering culms slender; bud initiation intravaginal or extravaginal. Leaf sheaths glabrous with membranous margins; blades filiform, pointed; ligules obtuse, membranous. Inflorescences paniculate, open; capillary branches dichotomously spreading, with higher order pedicels, minutely scabrous upward. Spikelets oblong, 2-flowered, purplish to silvery; glumes slightly unequal, lower glumes lanceolate, 1-nerved, upper glumes lanceolate, 3-nerved, often scaberulose on the midnerve and near apex; lemmas oblong, membranous, back rounded, 4(5)-toothed, teeth minute, central teeth larger than the lateral, rough; awn bent and twisted, inserted at the base or in the lower 1/3 of lemma; palea 2-keeled, shorter than the lemma, hyaline or sometimes membranous, scaberulose on the keels, apically erose; anthers 3. Caryopses ovoid, purplish; hilum punctiform. 1. Avenella flexuosa (L.) Drejer, Fl. Excurs. Hafn. 32. 1838. Basionym: Aira flexuosa L., Sp. Pl. 1: 65. 1753. Deschampsia flexuosa (L.) Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 2(1): 9. 1836, non Deschampsia flexuosa (L.) Nees, Gen. Fl. Germ. 2(1): t. 43. 1843, comb. superfl. Avenella flexuosa (L.) Parl., Fl. Ital. 1: 246. 1848, nom. superfl. Lerchenfeldia flexuosa (L.) Schur, Enum. Pl. Transsilv. 753. 1866. Podianapus flexuosa (L.) Dulac, Fl. Hautes-Pyrénées. 83. 1867. Salmasia flexuosa (L.) Bubani, Fl. Pyrene (Bubani): 4. 319. 1901. TYPE: Europe. ‘‘in petris, rupibus’’ (lectotype, designated by Clayton in MilneRedhead & Polhill [1970: 94], LINN 85.11 not seen, photo S!). Aira versicolor Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 2: 679. 1817. TYPE: Argentina. Islas Malvinas [Falkland Islands]: Portu Egmont., 1789, L. Neé s.n. (holotype, MA not seen; isotype, BAA!). Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:48 143 Cust # 2008115 144 Annals of the Missouri Botanical Garden Deschampsia tenella Phil., Anales Univ. Chile 94: 25. 1896, nom. illeg. hom., non Deschampsia tenella Petrie, Trans. & Proc. New Zealand Inst. 23: 402. 1891. Deschampsia philippii Macloskie, Rep. Princeton Univ. Exp. Patagonia, Botany 8: 961. 1906, as ‘‘philippi.’’ TYPE: Chile. ‘‘Valle fluminis Palena,’’ Jan. 1887, F. Delfin s.n. (holotype, SGO not seen; isotype, BAA fragm. ex SGO!). Aira vestita Steud., Syn. Pl. Glumac. 1: 424, tab. 49b. 1854. Deschampsia vestita (Steud.) Hauman, in Hauman & Parodi, Physis (Buenos Aires) 9: 337. 1929. TYPE: Chile. Sandy Point, s.d., W. Lechler 1193 (holotype, P!; isotypes, BAA ex P!, BAA ex S!, GOET not seen, fragm. US 02695871 not seen). Deschampsia martinii Phil., Anales Univ. Chile 94: 24. 1896. TYPE: Argentina. ‘‘Insulis Maclovianis,’’ Dec. 1884, Martin s.n. (holotype, SGO not seen; isotype, BAA ex SGO!). Deschampsia macloviana Gand., Bull. Soc. Bot. France 60: 28. 1913. TYPE: Argentina. ‘‘Falkland Islands, ad East F. Port Harriet,’’ s.d., C. Skottsberg 124 (holotype, S not seen; isotype, BAA!, fragm. ex S!). Discussion. Avenella flexuosa has been cited as introduced for Costa Rica (Davidse, 1994) in altitudes of ca. 3000 m. Its presence has been recorded outside the normal distribution range but in a suitable habitat. Aira flexuosa var. montana Brongn. (in Duperrey, Voy. Monde, Phan. 2: 23. 1829, nom. illeg. hom.) is a later homonym. Specimens from the Falkland Islands (Islas Malvinas) do not differ from continental specimens. Perennial, caespitose grass, new culms sometimes protruding through the base of the sheaths (extravaginal branching), sometimes rhizomatous; culms 35–70 cm tall, 2- or 3-noded, erect or bent, sheaths glabrous with membranous margins. Leaf blades 5.5–10 cm 3 0.5– 1 mm, conduplicate to filiform, stiff, pointed, glabrous adaxially, somewhat papillate abaxially; ligules 2.5– 3 mm, obtuse, membranous. Panicles 7.5–12.5 3 2– 5 cm, open and loose, with 5 to 7 verticils, branches spreading from the main fertile axis, dichotomously spreading, with higher-order pedicels, minutely scabrous upward. Spikelets oblong, 2-flowered, purplish to silvery; lower glumes 3.5–5 mm, lanceolate, 1-nerved, upper glumes 4.5–6 mm, lanceolate, 3-nerved, often scaberulose on the midnerve and near apex; lemmas 4– 5 mm, oblong, membranous, adaxially rounded, scabrid in the upper 1/3, 4(5)-toothed, teeth minute, central teeth larger than lateral teeth, rough; awn 5.5–7 mm, bent and twisted, inserted at the base or from the lower 1/3 of lemma; palea 3–5 mm, 2-keeled, hyaline or sometimes membranous, scaberulose on the keels, erose near the apex; anthers 1.5–2.5 mm. Caryopsis 1–2 mm, ovoid, brownish-purple. Chromosome number. 2n 5 28 (Hubbard, 1984, as Deschampsia flexuosa). Specimens examined. ARGENTINA. Chubut: Dpto. Futaleufú, región del Rı́o Corcovado, Cholila, 43uS, 71uW, N. Illı́n s.n. (CORD); Lago La Plata, A. Soriano 3137 (BAA); Dpto. Rı́o Senguerr, Lago Fontana, A. Castellanos 5946 (BA, S); Lomada del Coyte, fondo de valle, R. León 2364 (BAA). Santa Cruz: Dpto. Güer Aike, 3 km NW de Villa Minera Rı́o Turbio, S. Leuenberger & S. Arroyo 3667 (BAB); 26 km S of Rı́o Gallegos, W. J. Eyerdam et al. 24084 (SI); Dpto. Lago Argentino, Lago Argentino, P. James 40 (SI); Parque Nac. Los Glaciares, camping Arroyo Correntoso, 50u299140S, 72u579300W, A. Cocucci & A. Sérsic 2493 (CORD); Dpto. Rı́o Chico, alto Rı́o Blanco, R. N. Luti 3749 (CORD); Dpto. Deseado, Puerto Deseado, E. Ancibor & A. Vizinis 4421 (BAA). Tierra del Fuego: Dpto. Ushuaia, Ushuaia, R. N. Luti 1645 (CORD); Lapataia, S. Crespo s.n. (SI); Archipel d’Ushuaia, 25 Feb. 1896, N. Alboff s.n. (CORD); Ushuaia, 7 Apr. 1896, N. Alboff s.n. (CORD); Dpto. Rı́o Grande, near Rı́o Grande, ca. 1 m from ocean, Y. Mexı́a 7911 (S); San Sebastián, P. Dusén 304 (CORD); Rt. Nac. 3, San Sebastián, A. T. Hunziker 8266, 8275 (CORD); Islas Malvinas, Port Stanley, E. A. Ulibarri et al. 1073 (SI); Isles Malouines, D. Sellow s.n., Voyage D’Urville 1825 (P). CHILE. Aisén Region: Lago Buenos Aires, Valle León, I. von Rentzell 6280 (SI). Biobı́o Region: Lirquén, Quebrada Honda, K. Behn 20267 (CONC). Los Lagos Region: Llanquihue, Cerro Vichadero, Casa Pangue, A. Pfister 13568 (US). Magallanes and Antártica Chilena Region: Última Esperanza, Cord. Paine, M. T. K. Arroyo et al. 92-316 (CONC); Parque Nac. Torres del Paine, Cerro Diente, M. T. K. Arroyo & F. Squeo 85-0891 (CONC); Cerro Donoso, sect. Rı́o Las Chinas, M. T. K. Arroyo et al. 87-0219 (CONC); Punta Arenas, C. Montero 6206 (CONC); 78 km NW of Punta Arenas, rd. to Puerto Natales, W. J. Eyerdam et al. 24165 (SI); Isla Navarino, Puerto Williams, Cerro Bandera, F. Schlegel 8119 (CONC); Île Navarin, Sierres Rocalleuses, 25 Feb. 1896, N. Alboff s.n. (CORD); Punta Arenas, Chabunco, Pfister & Ricardi 11953 (BAA); Cerros del Club de Ski Punta Arenas, Pfister & Ricardi 11720 (BAA); Detroit de Magellan, Port Famine, Voyage de l’Astrolabe et de la Zeléc, 1838–1840, M. Jacquinot Hombron s.n. (BAA). Nicora (1978: 230). II. Vahlodea Fr., Bot. Not. 141, 178. 1842. TYPE: Vahlodea atropurpurea (Wahlenb.) Fr. ex Hartm. Distribution, habitat, and phenology. Avenella flexuosa is a common species that is distributed throughout Europe, eastern North America, northeastern Asia (Kamchatka to Japan), New Zealand, high mountains in west-central Africa (Kilimanjaro Mountains), and southern South America. It grows in humid soils, in open habitats or at the edge of temperate forests, from sea level to 1000 m elevation. Flowering occurs from December to March. Caespitose perennials; bud initiation often extravaginal. Leaf sheaths glabrous to rather scabrous, margins membranous or more rarely scarious; blades flat, pilose, middle nerve prominent with varying number of secondary nerves, pointed; ligules ovate to obtuse, membranous. Inflorescences paniculate, erect, open; branches lax, slender, nodding, scabrous. Spikelets ovoid, 2-flowered, mostly violaceous, pedicels scabrous; rachilla pilose, hairs about half the Illustration. Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:48 144 Cust # 2008115 Volume 97, Number 2 2010 Chiapella & Zuloaga Revision of Deschampsia, Avenella, and Vahlodea length of the lemma; glumes equal or slightly unequal, lower glumes broadly lanceolate, 1-nerved, keeled, upper glumes lanceolate, longer, 3-nerved, usually only the midnerve scabrous; lemmas 4(6)-toothed, teeth minute, equal, membranous; awns straight or geniculate, twisted, slightly exserted, stout, scabrous, inserted in the upper half of the lemma, rarely in the middle or lower; paleae hyaline, shorter than the lemma, 2-keeled; keels scabrous, sometimes slightly scabrous between the keels; anthers 3, purplish. Caryopsis fusiform. 5 mm, broadly lanceolate, 1-nerved, keeled, upper glumes 4.5–5.5 mm, lanceolate, 3-nerved, usually only the midnerve scabrous; lemmas 4(6)-toothed, teeth minute, equal, 2–4 mm, membranous, awns 2– 4 mm, usually geniculate or occasionally straight, twisted, slightly exserted, stout, scabrous, inserted in the upper half, rarely in the lower; paleae 2–3.5 mm, hyaline, 2-keeled, keels scabrous, sometimes slightly scabrous between the keels; anthers 0.8–1.5 mm, purplish. Caryopsis 0.5–1 mm, narrowly fusiform, brown to brownish red. Chromosome number. 1. Vahlodea atropurpurea (Wahlenb.) Fr. ex Hartm., Handb. Skand. Fl. (ed. 4): 30. 1843. Basionym: Aira atropurpurea Wahlenb., Fl. Lapp. (Wahlenberg): 37. 1812. Holcus atropurpureus (Wahlenb.) Wahlenb., Svensk Bot. Tidskr. pl. 687. 1826. Avena atropurpurea (Wahlenb.) Link, Hort. Berol. 1: 119. 1827. Deschampsia atropurpurea (Wahlenb.) Scheele, Flora 27: 56. 1844. TYPE: Finland. ‘‘Hab. in fruticetis campis et juniperetis subuliginosis per partem subsylvaticam totius Lapponiae passim copiose. . . in Enare juxta Jvalojoki et Sotajoki,’’ 22 Aug. 1802, G. Wahlenberg s.n. (lectotype, designated by Moberg & Nilsson [1991: 293], UPS not seen, photo ex UPS!; duplicates, LE-TRIN 1856.01a!, BAA!). Aira magellanica Hook. f., Fl. Antarct. 2: 376, t. 134. 1846. Aira atropurpurea var. magellanica (Hook. f.) Skottsb., Kongl. Svenska Vetensk.-Akad. Handl. 56(5): 174. 1916. Vahlodea atropurpurea subsp. magellanica (Hook. f.) Hyl., Bot. Not. 3: 356. 1953. Vahlodea magellanica (Hook. f.) Tzvelev, Bot. Mater. Gerb. Bot. Inst. Komarova Acad. Nauk SSSR 22: 68. 1963. TYPE: Chile. Port Famine, s.d., James Anderson King s.n. (holotype, K!; isotypes, BAA fragm. ex K!, photo US 867650 ex K!). Deschampsia brachyphylla Phil., Anales Univ. Chile 94: 23. 1896. TYPE: Chile. ‘‘Habitat in valle fluminis Palena,’’ 1887, Delfin s.n. (holotype, SGO 37212 not seen; isotype, BAA fragm. ex SGO!). Perennial, tufts loose, extravaginal or intravaginal shoots; culms 30–80 cm tall, 1- to 3-noded, the nodes brown. Leaf blades 2.5–6 cm 3 1–4 mm, flat, abaxially glabrous to sparsely pilose, adaxially pilose, trichomes soft, white, ca. 1 mm, sometimes abaxially missing in very old specimens, middle nerve prominent with 4 to 6 nerves on each side, pointed; ligules 3.5–5 mm, ovate to obtuse, membranous. Panicles 5– 8.5 3 1.5–4 cm, lax, open, branches slender, nodding, scabrous, more densely scabrous close to the spikelets. Spikelets 2-flowered, ovoid, 6 clustered at branch apices, green to violaceous, or gold, pedicels scabrous, rachilla villose, hairs reaching about half the length of the lemma; lower glumes 4– 145 2n 5 14 (Albers, 1972a, b). Illustration. Hitchcock et al. (1969: 548) (as Deschampsia atropurpurea). Distribution, habitat, and phenology. Vahlodea atropurpurea has a primarily Northern Hemisphere, circumpolar, amphi-atlantic distribution; in southern South America it is distributed from 32uS to 54uS (Strait of Magellan region), where it is found on rocky, humid slopes, between 400 and 2300 m elevation. Flowering occurs between January and March. Discussion. The populations in South America (vs. those in the Northern Hemisphere) have been treated as a different subspecies by Hultén (1941, 1968) and Haraldsen et al. (1991); the latter authors found that low genetic variation among Canadian and Norwegian populations was correlated with fewer morphological differences, thus giving little support for treatment as separate taxa. No molecular studies have been performed on South American populations. The morphological similarity of Northern and Southern Hemisphere plants is supported by characters reported in Haraldsen et al. (1991: 316, table 5): leaf length is slightly longer in South American plants; spikelet length and awn length are slightly shorter in South American plants. Treatment of the southern populations as a separate taxon is not warranted, and we think more detailed comparative morphological and molecular studies are needed to address this question. Specimens examined. ARGENTINA. Chubut: Dpto. Futaleufú, Parque Nac. Los Alerces, Lago Futalaufquen, orilla Este, A. Soriano 4169 (BAA). Mendoza: Dpto. San Rafael, valle del rı́o Atuel, mina de azufre, 80 km W de El Sosneado, O. Boelcke et al. 10213 (SI). Neuquén: Dpto. Minas, extremo N de la Laguna Varvarco Campos, O. Boelcke et al. 14217 (SI); Dpto. Huiliches, Volcán Lanı́n, M. N. Correa et al. 5688 (BAB); Dpto. Lácar, Parque Nac. Lanı́n, Lago Lácar, Cerro Malo, R. León & C. E. Calderón 1259 (BAA); Dpto. Los Lagos, Laguna Las Mellizas, valle Millaqueo, J. Diem 963 (SI); Cerro Colorado, J. Diem 253 (BAA); Parque Nac. Nahuel Huapi, Portal Pantojo, L. Cusato 2628 (BAA); nacimiento Arroyo Minero, O. Boelcke & M. N. Correa 7216 (BAA); Cerro 2u Mojón, O. Boelcke & M. N. Correa 7129 (BAA). Rı́o Negro: Dpto. Bariloche, Cerro López, L. R. Parodi 11506 (BAA); A. Burkart 6150 (BAB); I. Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:49 145 Cust # 2008115 146 Annals of the Missouri Botanical Garden von Rentzell 14661 (SI); Cerro Catedral, U. Eskuche 328 (BAA); Laguna Frı́as, camino a Cerro Riggi, R. Pérez Moreau s.n. (BA-34986, BAA); Cerro Riggi, A. L. Cabrera 6055 (BAA); Paso de Las Nubes, A. L. Cabrera 5915 (BAA). Santa Cruz: Dpto. Lago Argentino, Lago San Martı́n, desembocadura del Rı́o Fósiles, 19 Jan. 1918, A. Bonarelli s.n. (BA 39307). Tierra del Fuego: Dpto. Ushuaia, Sierra Valdivieso, paso Rı́o Azopardo, C. Skottsberg 240 (S); Cerca de Castillo, bosque de la Matanza, 31 Jan. 1942, A. Castellanos s.n. (BA 45566). CHILE. Aisén Region: Lago O’Higgins, Florida, A. Donat 527 (BAA); V. Thënmayer?, Feb. 1933, A. Donat s.n. (BAA). Magallanes and Antártica Chilena Region: Lago El Parrillar, E. Pisano V. 2519 (CONC); Cordillera del Paine, Jan. 1931, A. Donat s.n. (BAA); Penı́nsula Brunswick, A. Donat 455 (BAA). disarticulating above the glumes; rachillas prolonged beyond the upper floret, pubescent; glumes approximately as long as the spikelets, usually 1- to 3-nerved, keeled to rounded, usually covering the florets, roughly equal to unequal in size, membranous to scarious, apex acute; lemma 3- to 5-nerved, the central nerve usually continuing as the awn, awn dorsal, straight to bent, sometimes twisted below its lower half, inserted from the top to the base of the lemma, sometimes reduced to a minute appendage, lemma apex (2)4-toothed, thin; paleae generally as long as the lemma, 2-keeled, hyaline; lodicules 2, lanceolate to acute or lobed, frequently wider above the middle; ovary glabrate, with a few apical trichomes; anthers 3. Caryopsis ovoid to fusiform, when mature often fused with the palea; hilum elliptic; embryo small; endosperm hard to soft. III. Deschampsia P. Beauv., Ess. Agrostogr. 91–92, pl. 18, f. 3. 1812. TYPE: Deschampsia cespitosa (L.) P. Beauv. Monandraira E. Desv., Fl. Chil. (Gay) 6: 341. 1854. TYPE: Monandraira berteroana (Kunth) E. Desv. Airidium Steud., Syn. Pl. Glumac. 1: 423. 1854. TYPE: Airidium elegantulum Steud. Aristavena F. Albers & Butzin, Willdenowia 8(1): 83. 1977. TYPE: Aristavena setacea (Huds.) F. Albers & Butzin. Plants perennial or annual, often forming tussocks; culms erect, usually , 150 cm tall, sometimes slightly bent at the base, slender to stout. Leaf blades flat, folded, or convolute, glabrous or pubescent; ligules membranous. Inflorescences paniculate, open to contracted. Spikelets 2-flowered (rarely 1 or 3), usually perfect, sometimes cleistogamous, somewhat compressed, typically purple or violaceous to pale green, KEY TO THE SPECIES AND VARIETIES OF DESCHAMPSIA IN Discussion. Cheeseman (1906) noted that the awns of species of Deschampsia from New Zealand were extremely reduced and sometimes inserted close to the top of the lemma; this is a unique feature in the genus, which generally has awns that are developed and inserted on the lower 1/3 or near the base of the lemma. Parodi (1949) provided a description of the genus and considered this variation. The preliminary molecular data, available from nuclear ITS and plastid trnL sequences, support the inclusion of the New Zealand taxa in the genus. The two species D. chapmanii and D. tenella were included in the main clade of Deschampsia (Chiapella, 2007). SOUTH AMERICA 1a. Plants up to 50 cm. 2a. Plants annual. 3a. Leaf blades stiff, erect, linear, bristlelike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. D. airiformis 3b. Leaf blades flat to conduplicate, folded. 4a. Glumes with all dorsal nerves scabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. D. berteroana 4b. Glumes with only the dorsal midnerve scabrous. 5a. Awns 5–8 mm, inserted at the base of the lemma; panicles 5–25 cm . . . . . . 6. D. danthonioides 5b. Awns 7–10 mm, inserted in the middle of the lemma; panicles 4–12 cm . . . . . . 10. D. looseriana 2b. Plants perennial. 6a. Panicles 2–7.5 cm, slightly contracted to contracted in appearance; branches adpressed. 7a. Plants rhizomatous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. D. mendocina 7b. Plants caespitose. 8a. Awns stout, strongly bent or curved; spikelets 2- or 3-flowered . . . . . . . . . . . . . . 12. D. parvula 8b. Awns weak, straight, curved, or barely bent; spikelets 2-flowered. 9a. Callus hairs not reaching the middle of the lemma . . . 2. D. antarctica (Antarctic specimens) 9b. Callus hairs reaching the middle of the lemma, some hairs surpassing the middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. D. patula 6b. Panicles 3–18 cm, slightly contracted to an open aspect, branches not adpressed. 10a. Anthers pale yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. D. antarctica (continental specimens) 10b. Anthers pale orange to brownish red. 11a. Lemmas 3–4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. D. setacea 11b. Lemmas 2.5–3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. D. venustula 1b. Plants 50–125 cm (sometimes smaller in D. cespitosa, D. cordillerarum, D. elongata, and D. kingii). 12a. Panicles contracted, spikelike; both glumes 3-nerved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. D. elongata 12b. Panicles lax, open and wide; lower glume 1-nerved (exceptionally 3-nerved in D. cespitosa), upper glumes 3-nerved. 13a. Spikelets 3–5(–5.5) mm. 14a. Leaf blades with prominent adaxial ribs; nerves glabrous or slightly scabrid to sparsely scabrous. Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:49 146 Cust # 2008115 Volume 97, Number 2 2010 Chiapella & Zuloaga Revision of Deschampsia, Avenella, and Vahlodea 147 15a. Awns present in all florets, the awn inserted at the base or in the lower 1/3 of the lemma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4a. D. cespitosa var. cespitosa 15b. Awns often missing in the lower floret, the awn inserted in the superior 1/3 or near the apex of the lemma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4b. D. cespitosa var. pulchra 14b. Leaf blades without prominent adaxial ribs, nerves densely scabrous on both surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. D. cordillerarum 13b. Spikelets (4–)4.5–9.5 mm. 16a. Plants stout, densely caespitose; ligules acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. D. kingii 16b. Plants delicate, slender, not caespitose; ligules truncate . . . . . . . . . . . . . . . . . . . . . . . . 9. D. laxa The 15 species of Deschampsia described here are found mainly in the Andes of Argentina and Chile, from ca. 30uS to Tierra del Fuego, in a wide altitudinal range (300–4000 m), but mainly restricted to wet places near lakes, rivers, creeks, bogs, etc. Two localities outside this main distribution area with collections of D. cespitosa (in Bolivia and Brazil, see Specimens Examined) require further studies in order to determine whether they are introduced. No conservation threat has been detected for the species, although the high mountain habitats where some of them are found are particularly fragile. 1. Deschampsia airiformis (Steud.) Benth. & Hook. f., Gen. Pl. 3: 1158. 1883. Basionym: Trisetum airiforme Steud., Syn. Pl. Glumac. 1: 229. 1854. TYPE: Chile. ‘‘Arique, in pratis paludosis,’’ s.d., Herb. Lechler 723 (holotype, P!; isotypes, fragm. BAA ex P!, fragm. BAA ex K!, GOET not seen, K!, US 91466 not seen). Agrostis desvauxii Phil., Linnaea 33(3–4): 288. 1864. TYPE: Chile. ‘‘In andibus prov. Santiago,’’ Nov. 1861, R. A. Philippi s.n. (holotype, SGO PHIL-140 not seen, photo!; isotypes, BAA fragm. ex SGO!, K ex SGO photo!, SGO 37491!, US 556317 fragm. ex SGO PHIL-140 not seen). Annual with slender, delicate culms 4–15 cm tall, 1- or 2-noded, slightly bent at the base, sheaths glabrous with membranous margins. Leaf blades linear or narrowly linear to bristlelike, stiff, 1–6 cm 3 0.8–1.5 mm, adaxial side with the nerves moderately to densely scabrous, less dense abaxially; ligule elongate, 1.5–5.5 mm, scarious. Panicles contracted 1.5–5.5 3 0.5–2 cm, with 4 to 8 verticils, branches scabrous to densely hirsute. Spikelets (1)2flowered, green; rachilla with abundant pubescence in the upper half, glabrate or minutely pubescent below; lower glume narrowly lanceolate, 3–6 mm, (1 or 2)3nerved, upper glume narrowly lanceolate to lanceolate, 3–7 mm, 3-nerved, both glumes rough to minutely scabrous on the central nerve and distal 1/3, sparsely scaberulose between the nerves distally, margins scarious; lemma with apex 4(5)-toothed, lateral teeth acute and larger than the central ones; awn stout, bent, rarely straight and basally twisted, 3– 8 mm, inserted at the base or in the lower 1/3; palea hyaline, 1.5–3 mm, 2-keeled or rarely flattened, keels scabrous; anthers 1–2 mm, brownish red. Caryopsis 1–2 mm, ellipsoid, light brown. Illustration. Nicora (1978: 242). Distribution, habitat, and phenology. Deschampsia airiformis is found in the Andes of Argentina and Chile, from 40uS to 50uS latitude, between sea level and 800 m elevation. Flowering occurs in January and February. Discussion. Deschampsia airiformis was observed as often parasitized by the rust Tilletia cerebrina Ellis & Everh. (Nicora, 1978). This rust fungus (Ustilaginomycetes, Tilletiales) infects mostly pooid grasses as hosts (Castlebury et al., 2005). The distribution range of D. airiformis overlaps with that of D. berteroana, another annual of similar habit from central Chile, from which it differs in the shorter leaves and more contracted panicles with densely scabrous panicle branches. The overlapping and extended range of the annual species of Deschampsia agrees with Arroyo et al. (2002), who suggested that many elements of the high Andes flora might have used the high mountain corridor to faciliate migrations. Specimens examined. ARGENTINA. Chubut: Dpto. Tehuelches, Lago Cuatro, E. Nicora 9340 (SI); Rı́o Pico, Est. Tromenco, A. Soriano 4626 (BAB); Apeleg, Est. 3 Zorros, A. Soriano 5802 (BAA). Neuquén: Dpto. Minas, Laguna Varvarco Campos, arroyo Enfermera, 36u179S, 70u399W, O. Boelcke et al. 14056 (BAB); Dpto. Los Lagos, Est. Fortı́n Chacabuco, O. Boelcke 3201 (BAA); camino a Traful, L. R. Parodi 15358 (BAA). CHILE. Araucanı́a Region: Malleco, Lumaco, Santa Clara, G. Kunkel 631 (CONC). 2. Deschampsia antarctica E. Desv., Fl. Chil. (Gay) 6: 338. 1854. Replaced name: Aira antarctica Hook. f., Icon. Pl. 2: tab. 150. 1838, non Aira antarctica G. Forst., 1786. TYPE: Antarctica. New South Shetland Islands, s.d., Eights s.n. (holotype, K!). Airidium elegantulum Steud., Syn. Pl. Glumac. 1: 423. 1854. Deschampsia elegantula (Steud.) Parodi, Darwiniana 8: 452. 1949. TYPE: Chile. ‘‘Prope Punta Arenas,’’ Feb. 1853, W. Lechler 1220 (holotype, P!; isotypes, BAA! K!, US-76314 fragm. not seen). Deschampsia fuegina Phil., Anales Univ. Chile 94: 23. 1896. TYPE: Chile. Tierra del Fuego, Hab. Fuegia Orientalis, Feb. 1879, s. coll. (holotype, SGO PHIL-208 not seen, Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:50 147 Cust # 2008115 148 Annals of the Missouri Botanical Garden photo!; isotypes, BAA ex SGO!, US 556482 fragm. ex SGO PHIL-208 not seen). Deschampsia antarctica f. breviaristata Hack. in Dusén, Wiss. Ergebn. Schwed. Exped. Magellansl. 1895–1897 pt. 5: 221. 1900, nom. nud. the plants of Antarctica and some subantarctic islands, but typically open and exserted in continental individuals. It is possible, however, to find contracted panicles in Antarctic forms on the continent and plants with open, larger panicles in the subantarctic islands (the type specimen collected in the South Shetland Islands is a plant with an open, pyramidal panicle). The subtle morphological differences of both forms correspond to almost completely nonoverlapping areas of geographic distribution, making possible their differentiation as subspecies sensu Du Rietz (1930). The florets of the Antarctic plants are commonly cleistogamous, while plants with both chasmogamous and cleistogamous florets can be found in Tierra del Fuego. The molecular evidence available is limited, however, and has focused mainly on Antarctic and subantarctic island populations (Holderegger et al., 2003; Van de Vouw et al., 2007). A formal proposal of division in subspecies would require support of a study including several continental populations. Buschmann (1949) cited Deschampsia antarctica as introduced to Holland in 1936. The growth period of this species is particularly well suited for the rigorous conditions of Antarctica; the plants begin to grow during November, the flowering starts in the first week of January, and by the end of the month it is already possible to observe fertile spikelets. The quantity of fertile seeds produced, however, is smaller than those produced by plants growing outside Antarctica, e.g., in the Kerguelen Islands, Falkland Islands (Islas Malvinas), and Tierra del Fuego (Corte, 1961). Dusén (1900) published the name Deschampsia antarctica f. breviaristata, mentioning the specimen O. Nordenskjöld s.n. (Argentina, Patagonia australis: in valle superiore fluminis Gallegos), but because neither a valid description nor illustration is provided, it is considered a nomen nudum according to the International Code of Botanical Nomenclature (McNeill et al., 2006: Art. 32d). The search for the specimen collected by O. Nordenskjöld was not successful in the herbaria W (Hackel collection), S, and UPS. Perennial, forming dense caespitose tufts of basal leaves; culms slender, 10–25 cm tall, 1- or 2-noded, sheaths glabrous with margins membranous, more rarely scarious. Leaf blades conduplicate to filiform, 1.5–5.5 cm 3 1–1.5 mm, adaxial nerves scabrous, abaxially glabrous to minutely scabrous; ligules acuminate to acute, 2–9 mm, scarious. Panicles pyramidal, loose to contracted, when contracted partially included in the uppermost sheath, 4.5–15 3 1.5–6 cm, with 3 to 7 verticils; branches spreading, filiform, scabrous to densely hirsute at the distal 2/3. Spikelets 2(3)-flowered, green to violaceous, or variegated with both colors; lower glume narrowly lanceolate, 3–5.5 mm, 1-nerved, upper glume narrowly lanceolate to lanceolate, 4–6 mm, 3-nerved, both glumes ciliate on the central nerve and sometimes also scabrous between the nerves, basally glabrous, margins scarious; callus pilose, hairs short, not reaching midlemma in the non-antarctic specimens, even shorter in the antarctic specimens; rachilla pilose in the upper half; lemma bilobate, 4-toothed, lateral teeth longer than the central, scabrous to hirsute at the apex; awn straight, rarely geniculate, somewhat twisted at the base, 3–7 mm, weak, inserted in the lower 1/3, rarely at the middle, sometimes exserted, scabrous; palea hyaline, 2–3 mm, 2-keeled, keels moderately to densely scabrous; anthers 0.5–1.5 mm, pale yellow. Caryopsis 0.5–1.5 mm, ovoid, brown. Illustration. Hooker (1847: t. 133). Distribution, habitat, and phenology. Deschampsia antarctica is one of only two vascular plant species known to be native to Antarctica. The taxon is also found in several adjacent southern islands (Falkland Islands [Islas Malvinas], South Georgia, South Orkney, South Shetlands, Palmer Archipelago, Bouvet, Crozet, Kerguelen) and in the southern part of the American continent. Flowering occurs from November to February. Discussion. Deschampsia antarctica is a species similar to the Andean D. venustula and can be distinguished from this taxon by the longer awns that exceed the glumes and the spikelets, which are broadly lanceolate and purplish along the keels in D. venustula versus the spikelets being lanceolate and not colored in D. antarctica. The panicles in Deschampsia antarctica show clear dimorphism between the Antarctic and continental specimens, being closed, contracted, and often with the lower part included in the uppermost sheaths in Specimens examined. ARGENTINA. Chubut: Dpto. Tehuelches, Lago Vintter, E. Nicora 10231 (SI); Est. Caridad, 43u409S, 71u209W, costa del Rı́o Carrenleufú, A. Soriano 2554 (BAA, BAB); Valle Laguna Blanca, 71u159W, 45u529S, J. Koslowsky 224 (BAA); Valle Huenuleo, 45u559S, 71u509W, A. Soriano 3196½ (BAA); Barrancas a Corcovado, A. Soriano 5415 (BAA); Lago Vintter, A. Soriano 3083 (BAA); Alto Rı́o Senguerr, Est. Zootécnica Rı́o Mayo, A. Soriano 4474 (BAB); Dpto. Languiñeo, Tecka, Est. La Blanche, A. Soriano 2245 (BAB). Rı́o Negro: Dpto. Pilcaniyeu, Est. Rayhuau, O. Boelcke 4486 (BAB). Santa Cruz: Dpto. Lago Argentino, El Calafate, M. N. Correa et al. Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:50 148 Cust # 2008115 Volume 97, Number 2 2010 3082 (BAB); Dpto. Lago Buenos Aires, meseta del Lago Buenos Aires, Est. La Vizcaı́na, E. G. Nicora 26417 (BAA); Rı́o Coyle, Establ. Las Vegas, L. Dauber 168, 196 (BAB); Dpto. Güer Aike, Est. La Verdadera Argentina, 50u519S, 72u149W, s.d., S. Arroyo et al. s.n., T.B.P.A. 2139 (BAB); Laguna Cóndor, 51u469S, 71u409W, O. Boelcke et al. 12446 (BAB); prox. Puesto 2 Antonios, 51u379S, 72u119W, J. A. Ambrosetti & E. Méndez 206 (BAB); Dpto. Rı́o Chico, Lago Burmeister, Parque Nac. Perito Moreno, M. J. Dimitri & H. Correa Luna 8243 (BA). Tierra del Fuego: Dpto. Ushuaia, Ushuaia, N. Alboff 1027 (CORD), M. S. Pennington 392 (CORD); orillas del Canal Beagle, próx. base naval, M. N. Correa & R. L. Pérez Moreau 1923 (BAB); Est. Harberton, First Week Creek, D. M. Moore 1349 (BAB, K); Dpto. Rı́o Grande, ‘‘Fuegia Australis,’’ Rı́o Grande, 14 Jan. 1896, P. Dusén 405 (CORD); Est. Los Flamencos, 46 km W of Rı́o Grande, D. M. Moore & R. N. P. Goodall 276 (BAB, K); Rt. 3, Rı́o Grande, A. T. Hunziker 8225 (CORD); Golfo San Sebastián, Est. Sara, E. Grondona 4480 (BAA); Valle de Tierra Mayor, A. Ruiz Leal & F. A. Roig 15025 (BAA); Lago Fagnano, A. Castellanos 7572 (BAA); Laguna Grande, Sección Miranda, J. H. Hunziker 6763 (BAB); Antártida Argentina, Base Primavera, s.d., M. Laurı́a s.n. (BCRU); A. Corte 1 (BAA); Islas Biscoe, Tierra de Graham, F. Behn 12493 (BAA, S); Puerto Paraı́so, Cerro La Cruz, Z. Popovici s.n. (BA 67264); Islas Malvinas, Isla Soledad, Puerto Stanley, E. Ulibarri et al. 1067 (SI); Port Stephens, Cape Meredith, D. M. Moore 781 (K, S). Archipiélago Melchior: Isla Sobral, Punta Dos Monjes, costa SO, A. T. Hunziker 10190, 10196 (BAA, CORD); Isla Kappa, Punta NO, R. N. Luti 1485 (BAA, CORD); A. Martı́nez 2 (BAA). South Georgia Island: Stromness Bay, S. W. Greene 3058 (S, SI); Cumberland Bay, 15 May 1902, C. Skottsberg 63 (S). Palmer Archipelago: Pointe meridionale de l’île Anvers, 64u509S, 63u509W, 10 Nov. 1905, Dr. Turquets s.n., Exped. Antarctique Française (BAA, P); Île Anvers, baie de Biscue, 10 Feb. 1905, Dr. Turquets s.n. (CORD); Lysted Island, 64u199S, 62u539W, P. Siple 336 (BAA, P); Isla Sobral, costa NE, sobre el canal Murature, frente a Isla Piedrabuena, A. T. Hunziker 10200 (BAA, CORD). South Orkney Islands: Signy Island, W. J. L. Sladen H639/1 (BM). South Shetland Islands: King George Island, Marlet Inlet, Admiralty Bay STA. 1481 (BM); Caleta Potter, A. T. Hunziker 10147, 10150, 10152, 10153 (BAA, CORD); Isla Media Luna, peñasco meridional, A. T. Hunziker 10112 (CORD), promontorio rocoso al SO de la Isla, A. T. Hunziker 10121, 10205 (CORD); Isla 25 de Mayo, Base Jubany, s. coll. (CORD 1172). South Sandwich Islands: Candelmas Island, N of western lagoon, R. E. Longton 601 (BM). CHILE. Magallanes and Antártica Chilena Region: Isla Riesco, seno Skyring, Estancia Tita, A. Pfister & J. Ricardi 11938 (BAA); 15 km S of Punta Arenas, moisty sandy loam, W. J. Eyerdam et al. 24115 (K, S); 20 Feb. 1903, M. S. Pennington 187 (CORD); Última Esperanza, Lago Balmaceda, 51u539S, 72u159W, M. C. Latour et al. s.n., T.B.P.A. 1912 (BAB); Laguna Blanca, Feb. 1927, J. Guiñazú 183 (BAA); Cajon de Morales, 3000 m, Mar. 1931, F. Laffuel 1903 (BAA); inmediaciones de Punta Arenas y Rı́o de La Mina, 1 Mar. 1917, A. Bonarelli 96 (BAA); Curicó, Baños de Azufre del Planchón, 2700 m, Jan. 1933, G. Grandjot 15a (BAA); Cautı́n, A. Burkart 9507 (BAA). Kerguelen Islands: W. A. Procter 16 (BM); Kerguelén Station der Deutschen, 19 Feb. 1903, E. Werth s.n. (B); L’Aurore Australe, Lote 40, s.d., E. A. De La Rüe s.n. (B); 1908–1909, M. Bossieu s.n. (BAA). 3. Deschampsia berteroana (Kunth) Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Chiapella & Zuloaga Revision of Deschampsia, Avenella, and Vahlodea 149 Math., Seconde Pt. Sci. Nat. 4,2(1): 10. 1836. Basionym: Trisetum berteroanum Kunth, Révis. Gramin. 2: 457, tab. 142. 1831. Monandraira berteroana (Kunth) E. Desv., Fl. Chil. (Gay) 6: 343. 1854. Aira berteroniana (Kunth) Steud., Syn. Pl. Glumac. 1: 220. 1854. TYPE: Chile. Rancagua, Oct. 1828, C. L. G. Bertero 30 (lectotype, designated here, P!; duplicates, S!, SI ex P!, MO not seen, photo!). Deschampsia berteroana var. parvispicula Parodi, Darwiniana 8: 466. 1949. TYPE: Chile. Valparaiso, Limache, Cerro Cruz, A. Garaventa 1667½ (holotype, BAA!). Annual, slender, delicate, culms up to 45 cm tall, 1- to 3-noded, erect, sheaths glabrous with scarious margins. Leaf blades flat to rather conduplicate, folded, 4–11 cm 3 0.6–1.5 mm, nerves abaxially glabrous or with isolated prickles, scabrous adaxially; ligules acute, 3–7 mm, hyaline. Panicles loosely contracted, 5–20 3 2–2.5 cm, with 5 to 7 verticils, branches ascending, somewhat adpressed, scabrous to shortly pilose. Spikelets 2-flowered, erect and ascending, purplish green or purple variegated with green; lower glume narrowly lanceolate, 3.5–6 mm, 1- to 3-nerved, upper glume narrowly lanceolate to lanceolate, 4–6 mm, both glumes scabrous on all the nerves, rarely only the midnerve scabrous, glabrous or scabrous between the nerves, margins scarious; callus and rachilla pilose, trichomes of the callus short, not reaching the lower 1/3 of the lemma; lemma membranous, 3–4 mm, 4-toothed, lateral teeth longer than the central, (1)4-nerved; awn bent and twisted in the lower half, 6–9 mm, inserted at or near the lemma base, brown or dark brown in the lower part of the lemma, pale in the terminal portion; palea hyaline, 2– 3 mm, glabrous, bi-keeled, keels scabrous; anthers 1 to 3, 1.5–2 mm, pale yellow. Caryopsis 1–1.5 mm, fusiform, brownish red. Illustration. Parodi (1949: 465). Distribution and habitat. Deschampsia berteroana is endemic to the Andean region in central Chile and adjacent Argentina; it is found in open, humid soils, between 400 and 2800 m elevation. Although its presence has been recorded in several places in central Chile, it is considered vulnerable (Arancio et al., 2001). Discussion. Deschampsia berteroana belongs to a small group of four annual species with glumes with well-marked nerves, which also includes D. airiformis, D. looseriana, and D. danthonioides. The latter is shared with western North America, while the former are exclusive of the Andes between 30uS and 37uS. The high Andes of central Chile is a region with numerous annual species and a Mediterranean-type Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:50 149 Cust # 2008115 150 climate, consisting of wet winters followed by hot, dry summers (Arroyo et al., 1981). These conditions mostly favor short life cycles and the ability to seed before the winter. The species form an interesting group because of the annual habit, which might have developed as an adaptation to ecological constraints. Carlo Luigi Giuseppe Bertero (1789–1831) was an Italian botanist who drowned in a shipwreck while en route from Tahiti to Valparaı́so; he made collections in Chile and the Juan Fernández Islands, and most of his specimens were lost with him. Among the specimens left (found in P, S, SGO, and TO), the specimen Bertero 30 is a good example of Deschampsia berteroana and is therefore chosen as lectotype. Specimens examined. ARGENTINA. Mendoza: Las Heras, cerca de Polvaredas, 2210 m, J. A. Ambrosetti & E. Méndez 7713 (MERL); pasando Polvaredas, cerca del lugar de camping, 2200 m, J. A. Ambrosetti 1202 (MERL). CHILE. Araucanı́a Region: Temuco, termas de Rı́o Blanco, G. Kunkel 2004 (BAA). Biobı́o Region: Concepción, cerro del Seminario, E. Barros 3308 (BAA); Andes de Antuco, 1828, Dr. Pöppig s.n. (Hb. Trinius, LE); betw. Los Angeles & Santa Bárbara, T. Ryves & E. Clements 96/081 (K). Coquimbo Region: La Serena, E. Barros 1655 (CONC); La Rinconada, E. Barros 9913 (BAA); Fray Jorge, G. Martı́nez 52495 (CONC); Cordillera de Combarbalá, Potrero Grande, C. Jiles 6073 (CONC). Maule Region: Licantén, E. Barros 511 (BAA); Talca, Espinal de Los Llanos, O. Matthei & M. Quezada 1167 (CONC); Curicó, Cerro Condell, E. Barros 976 (BAA). Santiago Metropolitan Region: Santiago, s.d., R. A. Philippi s.n. (B); Apoquindo, G. Looser 1384 (BAA); Melipilla, H. Gunckel 20285 (CONC); Quebrada de Peñalolén, H. Gunckel 25067 (CONC); Batuco, H. Gunckel 26730 (CONC), H. Gunckel 18244 (S); Lo Prado, E. Barros 530 (BAA). Valparaı́so Region: Aconcagua, Jahuel, Santa Filomena, R. De Giorgio 452 (CONC); Limache, Cerro Cruz, A. Garaventa 1667 (BAA); El Sauzal, A. Garaventa 2838 (BAA). 4. Deschampsia cespitosa (L.) P. Beauv., Ess. Agrostogr. 91, 149, 160, pl. 18, fig. 3. 1812. Basionym: Aira caespitosa L., Sp. Pl.: 64. 1753, as ‘‘cespitosus.’’ Agrostis caespitosa (L.) Salisb., Prodr. Stirp. Chap. Allerton 25. 1796. Campella caespitosa (L.) Link, Hort. Berol. 1: 122. 1827. Avena caespitosa (L.) Kuntze, Taschen-Fl. Leipzig 45. 1867. Podionapus caespitosus (L.) Dulac, Fl. Hautes-Pyrénées 82. 1867. Aira major Syme subsp. caespitosa (L.) Syme ex Sowerby, Engl. Bot., ed. 3b: 11: 64. 1873. TYPE: ‘‘Habitat in Europae partis cultis & fertilibus’’ (lectotype, designated by Clayton, in Milne-Redhead & Polhill [1970: 92], LINN 85.8 not seen, photo!). 4a. Deschampsia cespitosa var. cespitosa. Deschampsia andina Phil., Anales Univ. Chile 43: 564. 1873. TYPE: Chile. Santiago, Valle del Yeso, Jan. 1866, R. A. Philippi s.n. (holotype, SGO PHIL-201 not Annals of the Missouri Botanical Garden seen, SGO photo!; isotypes, BAA ex SGO!, BAA fragm. ex K!, BAA 869 fragm. ex B!, SGO 37213 not seen, photo!, US 556497 ex SGO PHIL-201 not seen). Perennial, densely tufted grass with extremely variable habit, culms erect, 25–100(–120) cm tall, leafy at the base, sheaths glabrous with margins membranous to scarious. Leaf blades elongated, linear, flat or conduplicate, 8–20 cm 3 1.5–4(5) mm, adaxially scabrous, abaxially glabrous, with 6 to 8 prominent ribs, rough to the touch, margins often scarious; ligules obtuse to acute, 2–12 mm, membranous or scarious. Panicles lax, open, frequently nodding, 8–30 3 1.5–7 cm, rarely slightly contracted, peduncles glabrous or minutely scabrous. Spikelets (1)2(3)-flowered, compressed, purple with green and/ or gold (or a mix); rachilla usually with abundant hairiness, callus hairs short, about 1/3 the length of the lemma; glumes usually covering the florets, lower glume lanceolate, 3–4.5 mm, membranous to scarious, acute, entire, 1- to 3-nerved, upper glume narrowly elliptic, 3.5–5.5 mm, acute, 3-nerved, midnerve glabrous or scaberulose; lemma narrowly oblong, 2.5–3.5 mm, (3)4(5)-toothed to erose-toothed, lateral teeth larger, most rarely all equal or the central larger, (1)3- to 5-nerved, membranous; awn straight or slightly curved, weak, commonly not twisted, 1.5– 5.5 mm, inserted usually at the basal or median portions of the lemma and rarely exceeding the glumes, scabrous; palea hyaline, 2–3 mm, 2-keeled, keels scabrous. Anthers 1–2 mm, yellow or purple. Caryopsis 0.5–1 mm, ovoid, light brown. Chromosome number. Illustration. 2n 5 26 (Albers, 1980b). Nicora (1978: 230). Distribution, habitat, and phenology. This taxon has ca. 18 subspecies in Europe and Asia (Chiapella, 2000; Chiapella & Probatova, 2003) and six in North America (Chiapella et al., in prep.), which have an extensive synonymy and are not listed here. In South America, Deschampsia cespitosa can be found in wet meadows (called ‘‘mallines’’ in Patagonia), bogs, and along streams and creeks in the Andes from Bolivia to Tierra del Fuego. Isolated populations can also be found in high places of southeastern Brazi, at about 1000 m elevation. Flowering occurs between December and May. Discussion. Individual specimens of Deschampsia cespitosa from South America are similar in plant aspect, panicle, and spikelet size to those from Europe, although they are sometimes smaller. Although no specific study has addressed the origin of the South American populations, the limited molecular evidence available suggests differences between populations from both hemispheres, since northern Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:51 150 Cust # 2008115 Volume 97, Number 2 2010 and southern accessions were joined in separate clades (Chiapella, 2007). This species is similar in aspect to D. cordillerarum, from which it differs by having awns generally included in the glumes, paler spikelets, less densely scabrous leaves, and a more extensive geographic distribution and altitudinal range. While forms adjoined to D. cespitosa range from 300 to 4000 m, D. cordillerarum is found only above 1800 m. Specimens examined. ARGENTINA. Chubut: Dpto. Futaleufú, región del Rı́o Corcovado, entre El Bolsón y la Colonia 16 de Octubre, N. Illı́n 226 (CORD); Dpto. Rı́o Senguerr, Lago Fontana, Estancia Pepita, A. Soriano 1511 (SI); Valle de Gema, G. F. Gehrling 109 (SI); Lago Musters, Dec. 1939, E. Feruglio s.n. (BA 34811). La Rioja: Dpto. General Sarmiento, entre Laguna Brava y paso Pircas Negras, vega del Refugio Barrancas Blancas, F. Biurrun et al. 5280 (SI). Mendoza: Dpto. San Rafael, Valle del Rı́o Atuel, mina de azufre, W de El Sosneado, O. Boelcke et al. 10213 (SI); Dpto. San Carlos, Rı́o Diamante, Carette 22 (BA); Dpto. Malargüe, Valle Hermoso, J. R. Figueroa 3094 (CORD); 5 km W de Las Leñas, camino a Valle Hermoso, G. Seijo 1735 (BA); confluencia Rı́o Colorado con Arroyo Malalhue, A. Ruiz Leal 7843 (BA); Dpto. Tunuyán, camino al Real de Las Ovejas, 2830 m, A. Dalmasso 792 (MERL). Neuquén: Dpto. Minas, Laguna Varvarco Campos, Arroyo Benı́tez, 36u179S, 70u399W, O. Boelcke 14330 (SI); Dpto. Huiliches, Lago Tromen, Reynoso-Muller 3864 (BA); Paso Mamuil Malal, hito, R. A. Pérez Moreau s.n. (BA 37700); Pino Hachado, Feb. 1920, L. Hauman s.n. (BA 39287). Rı́o Negro: Dpto. Bariloche, pedreros del Tronador, M. J. Dimitri & H. Correa Luna 2843 (BA); Estancia El Cóndor, s.d., A. M. Faggi s.n. (BA 78632); Lago Guillelmo, 12 Feb. 1938, A. Castellanos s.n. (BA 21879); Dpto. Pilcaniyeu, orillas del Pilcañiú, camino Pilcaniyeu a Bariloche, E. Nicora 3768 (SI); Rı́o Ñirihuau, E de Bariloche, A. Burkart 6211 (SI); Los Juncos, 5 Mar. 1942, R. A. Pérez Moreau s.n. (BA 48435). San Juan: Dpto. Iglesia, Arroyo Tambillos, along trail Paso de Valeriano, 29u109S, 69u519W, I. M. Johnston 6096 (SGO); Dpto. Calingasta, Cordillera del Espinazito, Los Patillos, F. Kurtz 9667 (CORD); Valle Hermoso, F. Kurtz 9727 (CORD); Seitental, E. Ru 9695 (SI); Reserva Natural El Leoncito, Ciénaga de Los Cabreras, E. Haene 1979 (SI). BOLIVIA. Cochabamba: Chapare Province, trail betw. Laguna Corani & Rı́o Corani Mayu, N. Ritter & G. Crow 2199 (SI). BRAZIL. Paraná: General Carneiro, 20 km N of Iratim, L. Smith et al. 15713 (SI, US). Rı́o Grande do Sul: Cambará do Sul, Itaimbezinho, J. F. M. Valls 2394 (SI). Santa Catarina: Caçador, Rı́o Verde, 26u459S, 51u229W, L. Smith & R. Klein 13371 (SI). CHILE. Aisén Region: Coihaique Alto, M. Paz Martı́nez 7 (SGO); Paso Pehuenche, cerca de Lago Maule, M. J. Dimitri et al. 4455 (BA). Antofagasta Region: Vallenar, vic. of Laguna Chica, 28u489S, 69u529W, I. M. Johnston 5962 (S, SGO). Maule Region: Curicó, Cordillera Volcán Peteroa, E. Werdermann 582 (S, SI). Santiago Metropolitan Region: Santiago, Berg am Maipo tal bei San Gabriel, C. & G. Grandjot 3486a (BAA); Valle del Yeso, F. Schlegel 2590 (SGO). 4b. Deschampsia cespitosa var. pulchra (Nees & Meyen) Nicora, Darwiniana 18: 101. 1973. Basionym: Deschampsia pulchra Nees & Meyen, in Nees, Gramineae 24–25. 1841. Aira pulchra Chiapella & Zuloaga Revision of Deschampsia, Avenella, and Vahlodea 151 (Nees & Meyen) Steud., Syn. Pl. Glumac. 1: 220. 1854. TYPE: Chile. Cordillera de San Fernando, ad Rı́o Tinguiririca, 4000 m, Feb. 1831, F. J. F. Meyen s.n. (holotype, B!; isotypes, BAA fragm. ex P!, US 865601 fragm., photo ex B!). Plants 20–150 cm, panicles open. Leaf blades flat to folded, 10–30 cm; ligules acuminate, 5–10 mm. Spikelets 2-flowered, glumes equal, 3.5–5.5 mm, lower glume 1-nerved, upper glume 3-nerved; lemma 2.5–4 mm; awn straight, 0.5–2 mm, very weak, inserted in the superior 1/3 or near the apex, often lacking in the lower floret. Distribution and habitat. Deschampsia cespitosa var. pulchra is found in the Andes of Argentina and Chile, from San Juan Province to northern Neuquén Province, in humid mountain valleys. Discussion. This form is part of the extreme morphological variation of Deschampsia cespitosa and includes the plants rather smaller in height with reduced awns inserted in the upper 1/3 of the lemma. Some plants present normal 2-flowered spikelets, while others have the upper floret extremely reduced or absent, or have muticous florets. Specimens examined. ARGENTINA. Mendoza: Valle del Rı́o Atuel, O. Boelcke 4172 (BAB). Neuquén: Dpto. Ñorquı́n, termas de Copahue, A. L. Cabrera 6169 (BAA); Dpto. Minas, Cajon de Los Chenques, 36u289S, 70u489W, O. Boelcke et al. 13830 (BAB). San Juan: Yaguelito, F. Kurtz 9495a (CORD). CHILE. O’Higgins Region: Colchagua, Cordillera San Fernando, Rı́o Tinguiririca, s. coll. (BAA). 5. Deschampsia cordillerarum Hauman, Anales Soc. Ci. Argent. 86: 231, pl. 4. 1918. TYPE: Argentina. Mendoza: au bord de la riviere, a Las Cuevas, Mar. 1918, M. S. Pennington 22 (holotype, BA 39306!; isotypes, BAA ex BA!, US 865607 ex BA not seen). Perennial, densely caespitose, culms 35–60 cm tall, 1- or 2-noded, sheaths glabrous with membranous margins. Leaf blades folded, 5.5–12 cm 3 1–2 mm, acuminate, densely scabrous on the nerves on both sides, midrib not adaxially prominent; ligules acute, 5–11 mm, membranous, sometimes lacerate. Panicles loose, 8–13 3 2–6 cm, with 6 to 9 verticils, branches glabrous. Spikelets 2-flowered, violaceous to purplish, sometimes variegated with gold; lower glume narrowly lanceolate to lanceolate, 3–5 mm, 1-nerved, upper glume lanceolate, 4–5 mm, 3-nerved, scabrous on the midnerve, both glumes with margins membranous; lemma 3–4 mm, scarious to membranous, 4(5)toothed, lateral teeth longer than central tooth, 4(5)nerved, nerves glabrous; awn slightly bent and twisted in the lower half of the lemma, 5–6 mm, slender, Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:51 151 Cust # 2008115 152 Annals of the Missouri Botanical Garden inserted in the lower 1/3 or at the base, exceeding the glumes; palea narrow, 2–3 mm, 2-keeled, keels glabrous. Anthers 1–1.5 mm, yellowish to reddish. Caryopsis 0.3–0.8 mm, ovoid, brown. ligules acute, 3.5–7 mm, scarious. Panicles open, 5–25 3 4–8 cm, rather contracted in young plants, nodding and more open when mature, 6 or 7 verticils; branches often clustered in groups of 2, spreading from main axis, glabrous near the nodes, scabrous toward the distal portion. Spikelets 2-flowered, clustered toward the end of the branches; callus and rachilla pilose; lower glume narrowly lanceolate to lanceolate, 4–5 mm, 3-nerved, upper glume lanceolate, 4.5–6 mm, 3-nerved, glumes with the margins scarious and the nerves evident, scabrous normally only on the midnerve, rarely on all the nerves, and sometimes scaberulose between nerves; lemma 3.5–4.5 mm, 4-toothed, lateral teeth larger than central tooth, 4-nerved, sparsely scabrous on and between nerves, scarious; awn bent, twisted, 5–8 mm, scabrous, inserted at the base of the lemma. Anthers 0.5–1 mm, pale yellow to reddish. Caryopsis 1–1.5 mm, fusiform, brown. Illustration. Parodi (1949: 435). Distribution and habitat. Deschampsia cordillerarum is restricted to the Andes of central Argentina and neighboring areas of Chile, between 32uS and 35uS latitudes, at altitudes of 1800–3300 m, where it grows by small watercourses. Discussion. Deschampsia cordillerarum is similar to D. cespitosa in plant size and aspect, but differs in its slightly longer awns that exceed the glumes (vs. awns rarely exceeding the glumes in D. cespitosa), the more scabrid leaves, and the darker color of the spikelets varying between violaceous and purplish. Darker spikelets have been also noted for highaltitude forms of D. cespitosa in Europe (Vigo i Bonada, 1983). Specimens examined. ARGENTINA. Mendoza: Dpto. Las Heras, Las Cuevas, valle del rı́o Las Cuevas, arroyo afluente, 2 km del refugio militar Lamadrid, O. Boelcke et al. 9765 (BAA, SI). CHILE. Coquimbo Region: Ovalle, San Miguel, 30u509S, 70u359W, C. Jiles 3623 (CONC). Valparaı́so Region: Aconcagua, Laguna Castro, O. Zöllner 46212 (CONC). 6. Deschampsia danthonioides (Trin.) Munro, Pl. Hartw. 342. 1857. Basionym: Aira danthonioides Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1(1): 57. 1830. TYPE: ‘‘America Borealis Occidentalis,’’ 1829, Lindley s.n. (holotype, LE TRIN 1873.01c!). Deschampsia calycina J. Presl, Reliq. Haenk. 1: 251. 1830. Aira calycina (J. Presl) Steud., Syn. Pl. Glumac. 1: 220. 1854. TYPE: [Peru.] ‘‘Hab. In montanis Peruviae,’’ s.d., T. Haenke s.n. (holotype, PR not seen; isotypes, LE TRIN 1873.01a!, LE TRIN 1873.01b!, US 865608 photo ex PR!). Monandraira glauca E. Desv., Fl. Chil. (Gay) 6: 342. tab. 79, fig.1. 1854. Deschampsia glauca (E. Desv.) Parodi, Darwiniana 8: 467. 1949, non Deschampsia glauca Hartm., 1820. TYPE: Chile. ‘‘En lugares montañosos de la dehesa de Santiago,’’ s.d., C. Gay s.n. (holotype, P DESV 70!; isotypes, P!, US fragm. ex P DESV 70 not seen). Deschampsia gracilis Vasey, Bot. Gaz. 10: 224. 1885. TYPE: U.S.A. California: San Diego Co., 28 June 1884, C. R. Orcutt 1072 (holotype, PH not seen, photo!; isotypes, US 81797 not seen, photo!, GH not seen, photo!, MO not seen). Annual, slender, culms 15–35 cm tall, 1- to 2-noded, sheaths glabrous with membranous margins. Leaf blades flat to rather inrolled, narrowly lanceolate, 3.5–10 cm 3 0.8–1.5 mm, abaxial and adaxial sides both scabrous; Illustration. 263). Holmgren and Holmgren (1977: Distribution and habitat. Deschampsia danthonioides is an American species with a disjunct continental distribution. In western North America, it is found from Alaska to Baja California in wet or drying meadows, stream banks, or vernal pools; in South America, it occurs in north-central Chile from 29uS to 35uS latitudes in similar habitats. This species was introduced into England and Germany with seeds of Poa pratensis L. from the United States (Conert, 1987). Discussion. Deschampsia danthonioides differs from other Deschampsia species in the narrowly lanceolate spikelets and the panicle with scarce spikelets. The species is usually infected by the rust fungus Tilletia cerebrina (Ustilaginomycetes, Tilletiales) (Castlebury et al., 2005). Specimens examined. CANADA. British Columbia: Saltspring Island, near Isabelle Point S of Fulford Harbour, common in wet crevices of rock cliffs, J. A. Calder & K. T. MacKay 29623 (BM). U.S.A. California: San Gabriel Mtns., Horse Flats, V. Durán 3507 (P); Mendocino Co., Sherwood, A. S. Hitchcock 1419 (P); San Luis Obispo Co., 1 mi. from Creston, Shandon rd., R. F. Hoover 6789 (BAA); Lake Co., A. A. Beetle 1731 (BAA). Nevada: Elko Co., Pinon Range, foothill area just N of Cissillini Canyon, A. Tiehm & J. Nachlinger 14258 (CORD). Utah: Farmington, M. E. Jones 2157 (P). CHILE. Coquimbo Region: Coquimbo, al N de Mantos de Hornillos, 1 km antes de la Quebrada del Teniente, C. Marticorena & O. Matthei 143 (CONC); al pié de la cuesta de Buenos Aires, C. Marticorena & O. Matthei 228 (CONC); Elqui, cuesta de Buenos Aires, 29u349S, 71u149W, 500 m, C. Marticorena & O. Matthei 216 (B, CONC). O’Higgins Region: Rancagua, Termas de Cauquenes, 34u159S, 70u349W, A. Pfister 13102 (SI, US). Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:51 152 Cust # 2008115 Volume 97, Number 2 2010 Chiapella & Zuloaga Revision of Deschampsia, Avenella, and Vahlodea 7. Deschampsia elongata (Hook.) Munro, Pl. Hartw. 342. 1857. Basionym: Aira elongata Hook., Fl. Bor.-Amer. (Hooker) 2: 243, tab. 228. 1840. TYPE: U.S.A. Oregon: Sandy islands of the River Columbia, s.d., D. Douglas s.n. (holotype, K!; isotypes, BAA fragm. ex K!, US 76303 fragm. ex K photo!). airiformis is distinguished as a small, slender annual no higher than 15–20 cm, with panicle branches more scabrous to densely hirsute and stout awns to 8 mm long; D. elongata plants are perennial and range from 15–80 cm high, with shorter awns not exceeding 5 mm, and the panicle branches are sparsely scabrous. Aira aciphylla Franch., Miss. Sci. Cape Horn, Bot. 5: 384. 1889. Deschampsia aciphylla (Franch.) Speg., Anales Mus. Nac. Buenos Aires 5: 89. 1896. TYPE: Chile. Magallanes, Sep. 1877, L. Savatier 161 (holotype, P!; isotype, US 76296 fragm. ex P photo!). Aira aciphylla var. pumila Franch., Miss. Sci. Cape Horn, Bot. 5: 384. 1889. TYPE: Chile. Patagonie, Punta Arenas, 5 Feb. 1879, L. Savatier 196 (holotype, P!; isotypes, BAA fragm. ex P!, US 76297 fragm. ex P photo!). Perennial, caespitose, slender, culms densely tufted, 15–80 cm tall, 1- to 4-noded, with abundant innovations, sheaths glabrous with membranous margins. Leaf blades flat to rather folded, conduplicate, 4–12 cm 3 0.5–1.5 mm, glabrous abaxially, scabrous to densely scabrous adaxially, basal leaves often filiform, forming a dense cover; ligules acute, 3 to 10, scarious. Panicles contracted, spikelike, 8–30 3 0.5–2 cm, with 6 to 9 verticils, erect or slightly nodding, branches narrow, adpressed to the culm axis, sparsely scabrous. Spikelets 2-flowered; rachilla and callus pilose, with few long hairs ca. half the length of the lemma; lower glume narrowly lanceolate, 3.5– 4.5 mm, upper glume lanceolate, 3.5–5 mm, both glumes 3-nerved, violaceous or rarely green, shiny, scabrous on all nerves, most rarely between them, more densely scabrous in the distal portion of the lemma; lemma scarious, 2–3 mm, apex 4-toothed, lateral teeth larger than the central tooth; awn straight or nearly so, twisted, 2–5 mm, inserted between the middle and the base of the lemma, more often in the lower 1/3; palea hyaline, 1.5–2 mm, 2-keeled, scaberulose on the keels. Anthers 0.5–1.5 mm, reddish. Caryopsis 1–1.5 mm, fusiform, brown. Illustration. 261). Holmgren and Holmgren (1977: Distribution and habitat. Deschampsia elongata is a species with a disjunct distribution between western North America and South America. In North America, it is found along lake shores and timberline meadows from Alaska to California, whereas in South America it is commonly found in bogs, wetlands, and along small water courses in the Patagonian Andes of Argentina and Chile, from 30uS to 50uS latitudes. Discussion. Deschampsia airiformis and D. elongata are similar in appearance with their contracted, narrow panicles less than 2 cm wide. Deschampsia 153 Specimens examined. CANADA. British Columbia: Queen Charlotte Islands, Moresby Island, Gray Bay, common on sandy beach in clearings of woods near mouth of creek, J. A. Calder & R. Taylor 35255 (P). U.S.A. California: San Bernardino Co., cienaga betw. Bear Valley & Bluff Lake, L. Abrams 2833 (P); Truckee, A. S. Hitchcock 1417 (P). Idaho: Lake Waha, A. & E. Heller 3289 (P). Montana: Bozeman, J. W. Blankinship 599 (BM). Oregon: Curry Co., Rogue River, 4 mi. E of Gold Beach, D. Kildale 6100 (B). Washington: Olympic Mtn., A. Elmer 1664 (P); Cascade Mtns., upper valley of the Nesqually, O. D. Allen 38 (BM). ARGENTINA. Chubut: Dpto. Cushamen, Cholila, R. Martı́nez Crovetto 3028 (SI); Dpto. Futaleufú, Rı́o Corcovado, 71uW, 43uS, N. Illı́n 167 (CORD); región del Rı́o Corcovado, entre El Bolsón y Colonia 16 de Octubre, N. Illı́n 224 (CORD). Neuquén: Dpto. Huiliches, Parque Nac. Lanı́n, Volcán Lanı́n, Arroyo Rucu-Leufú, M. N. Correa et al. 5598 (BAB); Dpto. Los Lagos, Fortı́n Chacabuco, J. Vallerini 322 (SI); Lago Espejo, Parque Nac. Nahuel Huapi, E. Nicora 75317 (CORD). Rı́o Negro: Dpto. Pilcaniyeu, E. Nicora 3635 (SI); Dpto. Bariloche, Parque Nac. Nahuel Huapi, entre arroyo Apoco y lago Felipe, O. Boelcke & M. N. Correa 6051 (SI); Estancia El Cóndor, A. M. Faggi s.n. (BA 78782); Bariloche, L. R. Parodi 11425 (S). Santa Cruz: Dpto. Lago Argentino, 49u49S, 72u129W, s. coll. (SI 15053). CHILE. Aisén Region: Coihaique, E. Barros 5867 (K); Aisen, Km 49 del camino de Puerto Aisen a Coihaique, R. Maldonado 129 (B). Araucanı́a Region: Lonquimay, Cordillera Las Raı́ces, A. Burkart 9529 (SI); camino de Icalma a Liucura, en la estepa cerca de Marimenuco, cerca de la ribera de Bio Bı́o, A. Pfister 7383 (CONC). Magallanes and Antártica Chilena Region: Punta Arenas, Leña Dura, E. Barros 5871 (US); Patagonia Australis ad Punta Arenas, P. Dusén 549 (CORD); Rı́o El Ganso, seno Otway, R. Barrientos 220 (CONC); Última Esperanza, Cueva del Milodon, A. Ricardi & O. Matthei 375 (B, CONC); Puerto Natales, 11 Feb. 1936, E. Jara s.n. (CONC 71822); Lago Balmaceda, A. Kalela 2022 (S); Tierra del Fuego, Rı́o Condor, Forestal Trillium, Rı́o Calavera, E. Pisano et al. 8191 (CONC). MEXICO. Sierra de las Cruces, C. G. Pringle 4743 (P); Federal Distr., Eslava, C. G. Pringle 13244 (K). Jalisco: 15 mi. S of Autlán, near summits of mtns. below El Cuartón, R. McVaugh 10323 (BM); Cuautitlán, 2–3 km SE de Capillas, F. J. Santana Michel & L. Guzmán 3411 (US); Hidalgo, Pachuca, near Zerezo and below Parque Nac. El Chico, H. E. Moore Jr. 3126 (US). 8. Deschampsia kingii (Hook. f.) E. Desv., Fl. Chil. (Gay) 6: 335. 1854. Basionym: Aira kingii Hook. f., Fl. Antarct. 2: 376. tab. 135. 1846. TYPE: [Chile.] ‘‘South part of Tierra del Fuego, Strait of Magellan, Port Famine,’’ Jan. or Feb. 1833, C. Darwin 546 (lectotype, designated by D. M. Porter [1986: 30], K!; isotype, CGE not seen, photo!). Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:52 153 Cust # 2008115 154 Annals of the Missouri Botanical Garden Aira elatior Steud., Syn. Pl. Glumac. 1: 423. 1854. Trisetum dozei Franch., Miss. Sci. Cape Horn, Bot. 5: 384, pl. 9 & f. a–e. 1889, nom. superfl. TYPE: Chile. Sandy Point, W. Lechler 1222 (holotype, P!; isotypes, BAA fragm. ex P!, S!, US 76302 fragm. ex P, photo!). 58 km south of Punta Arenas at 58u369S, 70u559W (Ortiz-Troncoso, 1976). Type specimens of several species (Poa scaberula Hook. f., Trisetum cernuum Trin., and Geranium patagonicum Hook. f.) were collected in this locality, which was visited by many explorers, including Charles Darwin. Perennial, caespitose, sometimes rhizomatous, culms 25–125 cm tall, stout, erect, 1- to 3-noded, nodes glabrous, dark, sheaths striate with membranous margins, rarely scarious. Leaf blades flat to conduplicate, blades 4–15 cm 3 2–4.5 mm, acute, with scarious margins, nerves glabrous on the abaxial side, scabrous on the adaxial side to shortly pilose; ligules acuminate, 3.5–7 mm, membranous, sometimes lacerate. Panicles wide, open, pyramidal, 17–35 3 3–10 cm, with 5 to 10 verticils, branches scabrous. Spikelets 2(3)-flowered, purple to variegated with green or gold; callus and rachilla hairy, rarely glabrous; lower glume narrowly lanceolate, 4.5–9 mm, upper glume lanceolate, 5– 9.5 mm, both glumes with the dorsal nerve and 2 lateral nerves clearly marked at the base of glumes and disappearing toward the apex, nerves glabrous, margins membranous or scarious; lemma irregularly 4-toothed, teeth all similar, rarely the lateral or the central tooth larger; awn straight, strong, and short, twisted or not, 1– 4 mm, inserted in the middle or upper 1/3 of the lemma, less frequently in the lower 1/3; palea hyaline, becoming membranous when larger, 2–5 mm, 2-keeled, rarely flattened and scabrous between the keels, keels scabrous to densely hirsute; anthers 1.5–2.5 mm, reddish. Caryopsis fusiform, 1.5–2 mm, brown. Illustration. Hooker (1847: pl. 135). Distribution, habitat, and phenology. Deschampsia kingii is abundant in the understory of Nothofagus pumilio (Poepp. & Endl.) Reiche forests in southern Tierra del Fuego, where it also forms patches growing in water in swamps. It becomes less frequent to the north of Patagonia, where it is gradually replaced by D. cespitosa. The northern limit of its distribution is currently found at ca. 45uS. It can be found from sea level to 600 m elevation. Flowering occurs between January and March. Discussion. Deschampsia kingii is a stout plant similar to D. cespitosa, from which it differs by the larger spikelets with stronger awns and the denser pubescence on the leaves. The stands in Tierra del Fuego represent the typical form, while the differences with D. cespitosa become blurred in northern populations. The type locality mentioned in the protologue (Port Famine) is the name given by the English corsair Thomas Cavendish in 1587 to the settlement Rey Don Felipe, founded on 25 January 1584 by the Spanish Captain Pedro Sarmiento de Gamboa on the western side of the central part of the Strait of Magellan, ca. Specimens examined. ARGENTINA. Chubut: Dpto. Tehuelches, Valle Laguna Blanca, 45u529S, 71u159W, J. Koslowsky 142 (SI); Rı́o Pico, Estancia Tromencó, A. Soriano 5368 (BAB); Rı́o Aisen?, 1900, C. Burmeister s.n. (BAB); Cañadón de Gastre, T. Arneberg 9632 (BAB). Santa Cruz: Dpto. Lago Buenos Aires, entre Lago Buenos Aires N y Cabo Rı́o Mayer, F. Kurtz 32 (CORD); Dpto. Güer Aike, Estancia Stag River, afluente W del Rı́o Venados, meseta Latorre, 51u349S, 71u579W, O. Boelcke et al. TBPA 3284 (BAB); Dpto. Lago Argentino, Lago San Martı́n, brazo sud, Mar. 1933, L. R. Parodi s.n. (BAA). Tierra del Fuego: Monte Olivia, M. Awschalom s.n., Herb. Parodi 9550 (K, S); cerros próximos al Monte Olivia, R. N. Luti 1435 (CORD); Monte Olivia, A. T. Hunziker 8218 (CORD); Lago Fagnano, J. H. Hunziker 6709 (BAB); Estancia Moat, W. Bank, D. M. Moore 1688 (K); Ushuaia, M. S. Pennington 267 (CORD); Ladera Martiales, Feb. 1942, L. A. Tortorelli s.n. (BAB); Tierra Mayor Valley, R. N. P. Goodall 4700 (BAB); Isla de Los Estados, Caleta Brent, A. Castellanos 12846 (BA). CHILE. Magallanes and Antártica Chilena Region: Punta Arenas, Tres Brazos, F. Pastore 72 (SI); Patagonia Australis ad Punta Arenas, P. Dusén 548 (CORD); Istmo de Ofqui, expedición argentina ‘‘Hicken-Reichert’’ (SI 10985); Última Esperanza, puerto Bella Vista, 51u039S, 73u159W, F. Roig et al. s.n., TBPA 5472 (BAB); Lago Balmaceda, A. Kalela 2021 (S); 15 km S of Punta Arenas, W. J. Eyerdam et al. 24132 (S); Isla Navarino, C. Skottsberg s.n. (S); mina Loretto, Y. Mexı́a 7978 (K); Penı́nsula Taitao, San Rafael, M. Gusinde 474 (S); Isla Otaries, Surgidero Romanche, 55u379S, 67u329W, E. Pisano 5089 (SI). 9. Deschampsia laxa Phil., Linnaea 29: 92. 1858. TYPE: Chile. Chonos, en las playas de Guaytecas, R. Fonck 53 (holotype, SGO PHIL 199 not seen; isotypes, BAA ex SGO!, K photo ex SGO!, US 556490 fragm. ex SGO not seen). Calamagrostis hirthii Phil., Anales Univ. Chile 94: 22. 1896. TYPE: Chile. ‘‘In valle fluminis Palena,’’ Jan. 1885, A. Hirth s.n. (holotype, SGO PHIL 133 not seen; isotypes, BAA fragm. ex SGO!, SI photo ex SGO!, US 1939357 fragm. ex SGO not seen). Perennial, delicate, culms 50–90 cm tall, 1- to 4noded, sheaths glabrous with margins membranous to scarious. Leaf blades linear, conduplicate, 6–12 cm 3 1–4 mm, nerves glabrous on the abaxial side, rarely scabrous, on the adaxial side scabrous; ligules truncate, 4–10 mm, scarious, when larger membranous, basally dilated and prolonged on both sides of the leaf blade. Panicles loose, open, 15–25 3 3– 15 cm, with 6 to 11 verticils, panicle branches up to 12 cm, scabrous, grouped in the base of the panicles forming clusters of 3 to 5 branches, base slightly swollen. Spikelets 2-flowered, green to whitish, Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:52 154 Cust # 2008115 Volume 97, Number 2 2010 Chiapella & Zuloaga Revision of Deschampsia, Avenella, and Vahlodea sometimes variegated with gold; callus and rachilla hairy, hairiness less dense toward the distal end; glumes linear-lanceolate, lower glume 3.5–8 mm, 1-nerved, scarious on margins and toward the apex, upper glume 4–9 mm, 3-nerved, normally only the midnerve scabrous, margins membranous; lemma 2– 4 mm, membranous, 4(5)-toothed, lateral teeth slightly larger (rarely smaller) than the central tooth, 4(5 or 6)nerved, the nerves glabrous or sparsely scabrous in the distal portion; awn slightly bent, somewhat twisted in the base of the lemma, 3–7 mm, inserted adaxially in the lower to middle 1/3 of the lemma; palea 2-keeled or rarely rounded, 2–3 mm, hyaline to membranous, keels scabrous. Anthers 1.5–2 mm, reddish. Caryopsis narrowly fusiform, 1–1.5 mm, brown to reddish. scarious. Leaf blades flat or folded, 2.5–11 cm 3 0.5– 1.5 mm, nerves glabrous and somewhat papillate on the abaxial side, scabrous on the adaxial side; ligules acuminate, 3–7 mm, membranous. Panicles open to slightly contracted, 4–12 3 1–4 cm, erect or nodding, with 4 to 7 verticils, lower branches adpressed and grouped in clusters of (2)3, moderately to densely scabrous. Spikelets 2-flowered, pedicels scabrous; rachilla and callus of florets with white hairiness; glumes narrowly lanceolate, lower glume 5.5–7 mm, upper glume 6–7.5 mm, both 3-nerved, commonly with only the midnerve scabrous, with margins hyaline and lateral nerves vanishing toward the apex; lemma contracted in the upper half, 4–6 mm, minutely scabrous or papillate toward the apex, unequally 4-toothed, the lateral teeth are prolongations of the lateral nerves, central teeth minute, rudimentary; awn bent, twisted in the lower half, 7–10 mm, inserted in the middle of the lemma; palea 2-keeled, hyaline, scaberulose on the keels. Anthers 1 to 3, 0.5–2 mm, pale yellow. Caryopsis 0.5–1.5 mm, ovoidal to fusiform, brown to reddish. Illustration. Nicora (1978: 230). Distribution. Deschampsia laxa occurs in the southern Andes of Argentina and Chile, from 43uS (Rı́o Palena) to Tierra del Fuego; it occupies roughly the same areas as D. kingii, although it is much less abundant than that species. Discussion. Deschampsia laxa differs from D. kingii in the leaves, which are glabrous on the adaxial side; the ligules, which are truncate and prolonged into the sides of the leaf blade; the smaller spikelets; and the point of insertion of the awns into the lemmas (nearly basal in D. laxa vs. inserted in the middle of the lemma in D. kingii). The branches of the panicle are also flexuous and curved; otherwise, the two species are similar in aspect, and the identity of D. laxa has yet to be confirmed with more collections and molecular studies. Specimens examined. ARGENTINA. Chubut: Dpto. Cushamen, Esperanza Lake, Horqueta, A. E. Johnson 586 (SI); Dpto. Futaleufú, Parque Nac. Los Alerces, Lago Menéndez, R. Pérez Moreau s.n. (BA 49492); Futaleufú, A. Soriano 3493 (SI); ‘‘Patagonia,’’ s. loc., A. Bonarelli 78, Plantae Magellanicae (SI). CHILE. Magallanes and Antártica Chilena Region: Última Esperanza, Puerto Bella Vista, 51u309S, 73u159W, F. Roig et al. TBPA 5103 (BAB); Isla Hoste, caleta Awaiakirrh, E. Pisano 5474 (SI). 10. Deschampsia looseriana Parodi, Darwiniana 8: 460. 1949. TYPE: Chile. Santiago, Batuco, 500 m, 17 Sep. 1936, G. Looser 3439 (holotype, BAA!). Deschampsia looseriana var. triandra Parodi, Darwiniana 8: 464. 1949. TYPE: Chile. Santiago, Batuco, 26 Sep. 1931, G. Looser 2049 (holotype, BAA!). Annual, delicate, slender, culms 15–40 cm tall, 2to 3-noded, branched from the base, sheaths striate, glabrous, with margins membranous or less commonly Illustration. 155 Parodi (1949: 462). Distribution, habitat, and phenology. Deschampsia looseriana is found in central Chile, between Valparaiso and Curicó. It grows between 300 and 1200 m elevation. Flowering occurs between September and November. Discussion. Deschampsia looseriana and the other annuals of central Chile and adjacent regions of Argentina (D. berteroana and D. danthonioides) are poorly known species that have been evaluated as vulnerable (Arancio et al., 2001). Deschampsia looseriana var. triandra was established by Parodi (1949) based on the presence of flowers with three stamens, differing from the typical form with commonly one or two stamens. Modifications in stamen number, anther size, and other reproductive structures (e.g., reductions in lodicule and awn size, filament length) are typical features of cleistogamy (Campbell et al., 1983) and therefore are not considered as valid for separating a variety. Careful examination of the type of variety triandra (G. Looser 2049, BAA) shows the existence of flowers with one stamen with very reduced anthers, a fact already noted by Parodi (1949: 464). Specimens examined. CHILE. Biobı́o Region: Concepción, E. Barros 11 (CONC). Coquimbo Region: Coquimbo, Combarbalá, Cuesta de Punitaqui, C. Marticorena & O. Matthei 387 (CONC). Maule Region: Vichuquén, E. Barros 1637 (CONC); Constitución, Los Molinos, A. Barnier 232 (CONC). Santiago Metropolitan Region: Batuco, G. Montero 2366 (BAA); Batuco, G. Looser 3438 (CONC); La Reina, E. Navas 8123 (CONC); Quebrada de Peñalolén, Y. Bravo 23 (CONC); Maipú, Cerro El Águila, 32u549660S, 62u889140W, A. Tomé s.n. (CORD 1133). Valparaı́so Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:52 155 Cust # 2008115 156 Annals of the Missouri Botanical Garden Region: Valparaı́so, Marga Marga, A. Laffuel & B. Pirión 1838 (BAA); La Rinconada, U. Levi 2674 (CONC); Limache, Pangal, G. Looser 3718 (BAA). Densely caespitose perennial, culms 5–25 cm tall, 1-noded, sheaths glabrous, widened at the base, with membranous or scarious margins. Leaf blades linear to filiform, abundant, forming dense clumps, blades 1.5– 6 cm 3 0.5–1 mm, nerves glabrous on the abaxial side, sparsely scabrous on the adaxial side; ligules acuminate, 3–9 mm, scarious. Panicles contracted, lower part included in the uppermost sheath or slightly exserted, 3–7 3 1–2 cm, with 4 to 9 verticils, branches normally adpressed to the culm axis, densely scabrous. Spikelets 2(3)-flowered, erect, violaceous, often variegated with gold or green; callus and rachilla pilose, trichomes short and scarce; lower glume narrowly lanceolate to lanceolate, 3–7 mm, 1-nerved, upper glume 3.5–7.5 mm, 3-nerved, both glumes glabrous, normally scabrous only along the midnerve in the distal portion, with margins membranous or scarious; lemma 4-toothed, lateral teeth acute, larger than the central tooth, (3)4(5)-nerved, nerves glabrous; awns stout, exserted, strongly bent or curved, 3–6.5 mm, twisted in the lower half, inserted in the lower 1/3 of the lemma; palea hyaline, 2–3 mm, bikeeled, scabrous along the keels. Anthers 0.5–1 mm, reddish to pale orange. Caryopsis 0.5–1 mm, fusiform. 11. Deschampsia mendocina Parodi, Darwiniana 8: 447. 1949. TYPE: Argentina. Mendoza: Sierra de la Medialuna, valle, 1700 m, 15 Feb. 1922, C. Rigal s.n. (holotype, BAA 4685!). Perennial, rhizomatous, culms erect, slightly bent at the base, 20–25 cm tall, 2- to 3-noded, sheaths glabrous with margins scarious. Leaf blades folded, conduplicate, 6–8 cm 3 0.5–1 mm, nerves glabrous on the abaxial side, scabrous on the adaxial side, prickles and very short hairs mostly on the nerves; ligules acute, 4–6 mm, scarious. Panicles slightly contracted, subspiciform, 6–7.5 3 1–1.5 cm, peduncles scabrous, short. Spikelets 2- to 3-flowered, purplish, callus densely pubescent, trichomes white, reaching the top of the florets, rachilla pilose; lower glume 3.5–3.8 mm, 1-nerved, upper glume 3.5–4 mm, 3-nerved, both glumes with nerves glabrous and margins membranous, sometimes the middle nerve with isolated prickles; lemma 4-toothed, teeth similar or the central tooth slightly longer, 2–3 mm, 4- to 5-nerved, nerves glabrous; awn straight, not twisted, 1.5–3 mm, scabrous, inserted in the lower half or at the base of the lemma, not exceeding the glumes; palea 2-keeled, 2–3 mm, keels scabrous, hyaline. Anthers 0.5–1 mm, reddish-orange. Caryopsis not seen. Illustration. Parodi (1949: 448–449). Distribution and habitat. Deschampsia mendocina is a rare species known only from the type in the Andes of central Argentina. Discussion. Deschampsia mendocina is similar to D. cespitosa, but differs by its more compact aspect (culms only to 25 cm vs. to 100(120) cm in D. cespitosa), its smaller and more contracted panicles (only to 7.5 cm in vs. lax, open panicles up to 30 cm in D. cespitosa), and by the presence of rhizomes (vs. absent in D. cespitosa). 12. Deschampsia parvula (Hook. f.) E. Desv., Fl. Chil. (Gay) 6: 339. 1854. Basionym: Aira parvula Hook. f., Fl. Antarct. 2: 377. 1846. Trisetum parvulum (Hook. f.) Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 5: 89. 1896. Deschampsia parvula (Hook. f.) Macloskie, Rep. Princeton Univ. Exp. Patagonia, Botany, Volume viii, 1 [2], Botany 8(1,5,1): 202–203. 1904, nom. illeg. TYPE: Chile. Cape Horn, Hermitte Island, J. D. Hooker 12 (holotype, K!; isotype, BAA fragm. ex K!). Illustration. Nicora (1978: 234). Distribution, habitat, and phenology. Deschampsia parvula ranges from the Andes of southern Argentina and Chile, from Lago Argentino to Tierra del Fuego, occurring in wet bogs to rocky soils from sea level to 2800 m. Flowering occurs in January and February. Discussion. The contracted panicles of Deschampsia parvula are similar to the inflorescences of Trisetum, but the species clearly differs from the latter genus by the 4-toothed lemmas. Deschampsia parvula is similar to D. patula in the small plant size and the short leaves, but it differs from D. patula in its panicles, which are more contracted and with the branches close to the axis. Specimens examined. ARGENTINA. Santa Cruz: Dpto. Güer Aike, valle superior del rı́o Turbio, faldeo Cordillera Chica, 51u279S, 72u069W, J. Ambrosetti & E. Mendez, TBPA 3647 (BAB); cerca del ventisquero Moreno, F. Reichert et al. s.n., Iter Patagonicum 126 (SI). Tierra del Fuego: Dpto. Ushuaia, Cerro Redondo, E. Grondona 7226 (BAA); Sierra Lucas Bridges, Monte Spion Kop, D. M. Moore 2800 (K); Lashifashaj Valley, NE side, near Mt. Cornu, D. M. Moore 1758 (BAA); Estancia Moat, mtn. at head of Rı́o Chico, 54u539S, 66u539W, D. M. Moore 1654 (BAB); Monteurs audessus d’Ushuaia, rochers 800–900 m, 9 Feb. 1896, N. Alboff s.n. (CORD); Hauteurs de la rive droite in torrent Ushuaia, 29 Feb. 1896, N. Alboff s.n. (CORD); Isla de Los Estados, Puerto Abrigado, A. Castellanos 12831 (BA); Basket Island, C. Spegazzini 20701 (BAA). CHILE. Magallanes and Antártica Chilena Region: Última Esperanza, Penı́nsula Roca, Seno Resi, TBPA 2916 (SI); Punta Arenas, O. Zollner 9623 (CONC); Tierra del Fuego, Sector Vicuña, Forestal Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:53 156 Cust # 2008115 Volume 97, Number 2 2010 Chiapella & Zuloaga Revision of Deschampsia, Avenella, and Vahlodea Trillium, E. Pisano et al. 7577 (CONC); Parque Nac. Torres del Paine, Cerro Agudo, M. T .K. Arroyo & F. Squeo 87-0007 (CONC). Specimens examined. ARGENTINA. Chubut: Dpto. Rı́o Senguerr, Laguna Blanca, C. Burmeister s.n. (SI 15043); Dpto. Rı́o Senguerr, El Coyte, con Festuca argentina o F. pallescens, R. León 2400 (BAA). Santa Cruz: Dpto. Guer Aike, Laguna Cóndor, O. Boelcke 12446 (BAB); RN 3, 10 km W de Rı́o Gallegos, a orillas del rı́o Gallegos, O. Boelcke 12358 (BAB); sección San Antonio, 51u249S, 71u349W, F. Roig et al. 77 (SI). Tierra del Fuego: Dpto. Rı́o Grande, San Sebastián, Estancia José Menéndez, J. Vallerini 3900 (CRP); Rı́o Grande, Estancia M. Behety, M. Collantes 2107 (SI); Orange Harbor, U.S. South Pacific Exploring Expedition, US 867656 (US). CHILE. Magallanes and Antártica Chilena Region: Última Esperanza, Sierra de los Baguales, Cerro Santa Lucı́a, 50u449S, 72u209W, M. T. K. Arroyo 85155 (CONC); Isla Wollaston, Caleta Lientur, 55u449S, 67u199W, E. Pisano 5123 (SI). 13. Deschampsia patula (Phil.) Pilg. ex Skottsb., Kongl. Svenska Vetensk. Acad. Handl. 56: 175. 1916. Basionym: Monandraira patula Phil., Anales Univ. Chile 43: 565. 1873. Deschampsia elegantula var. patula (Phil.) Parodi, Darwiniana 8: 454. 1949. TYPE: Chile. Magallanes, Punta Arenas, 1869, s. coll. (holotype, SGO-PHIL 244 not seen, photo!; isotypes, SGO 37214, photo!, US 867651 fragm. ex SGO not seen, K photo ex SGO!). Perennial, caespitose, erect, culms 8–20 cm tall, 4- to 6-noded, sheaths glabrous with membranous margins. Leaf blades folded, 1.5–3.5 cm 3 0.5– 1.5 mm, abaxial side sparse to densely scabrous, prickles mainly on nerves, on the adaxial side prickles less abundant; ligules acute, sometimes lacerate, 2– 5 mm, scarious. Panicles open to contracted, 2.5– 6(10) 3 2–6 cm, with 4 to 6 verticils, branches moderately to densely scabrous. Spikelets 2-flowered, purple to gold, sometimes variegated with green; lower glume lanceolate, rarely narrowly lanceolate, 2.5– 6 mm, 1(3)-nerved, upper glume lanceolate, 2.5–7 mm, 1(3)-nerved, both glumes scabrous along the midnerve with margins scarious; callus pilose, hairs reaching the middle of the lemma, sometimes few hairs exceeding the middle; lemma 2–4 mm, 4-toothed, lateral teeth larger, rarely all similar, 1- to 4-nerved, nerves glabrous; awns weak, bent and normally twisted and not exceeding the glumes, inserted in the middle or lower 1/3 of the lemma, 1.5–6 mm, scabrous; palea 2-keeled or flattened, 1.5–4 mm, keels scabrous. Anthers 0.5–1 mm, dark reddish to violaceous. Caryopsis 0.5–1 mm, ovoid. Illustration. Nicora (1978: 234). Distribution and habitat. Deschampsia patula is found in the Andes of Argentina and Chile, from Santiago to Tierra del Fuego and also in the southern Patagonian steppe, primarily in wet bogs. It is found between 900 and 3500 m elevation. Discussion. Deschampsia patula is similar to D. antarctica, from which it differs mainly by its shorter, twisted awns, the awns not exceeding the glumes, the more contracted panicles, and the longer hairs of the callus. The existence of intermediate individuals between both taxa was noted by Parodi (1949), who described D. elegantula var. patula, probably trying to accommodate individuals of D. elegantula (5 D. antarctica) approaching D. patula. 157 14. Deschampsia setacea (Huds.) Hack., Cat. Rais. Gramin. Portugal 33. 1880. Basionym: Aira setacea Huds., Fl. Angl. (Hudson): 30. 1762. Aira montana var. setacea (Huds.) Huds., Fl. Angl., ed. 2: 35. 1778. Aristavena setacea (Huds.) F. Albers & Butzin, Willdenowia 8: 83. 1977. TYPE: United Kingdom. Litcham Common, 18 July 1883, F. J. Hanbury s.n. (neotype, designated by Chiapella [2009: 242], BM not seen, photo!). Perennial, caespitose, with vegetative shoots densely packed, erect, culms 12–45 cm tall, 2- to 3-noded, sheaths glabrous with membranous margins. Leaf blades bristlelike, blades 2.5–10 cm 3 1–1.5 mm, inrolled, sharply pointed, glabrous to sparsely scabrous on the abaxial side, margin of blade scabrous, with abundant prickles, adaxial side scabrous, prickles more abundant on nerves; ligules narrowly lanceolate, 4.5–11 mm, hyaline, acuminate. Panicles loose, lanceolate, 8–18 3 1.5–5 cm, with 5 to 7 verticils, main axis glabrous to scaberulose, lower branches verticillate, branches sparsely scabrous, brown to purplish, darker toward the tips. Spikelets 2-flowered, purplish, often clustered at the end of branches; lower glume narrowly lanceolate, (3–)4–5 mm, 1-nerved, upper glumes lanceolate (3.5–)4.5–6 mm, 3-nerved, both glumes membranous, glabrous to sparsely scabrous in the nerves; callus pilose, hairs short, 6 adpressed to the rachilla, lemma 3–4 mm, 4(5)-toothed, teeth irregular, the lateral longer; awn bent and twisted, 4.5–6 mm, inserted in the inferior 1/3 or at the base, brownish in the abaxial half, purple in the adaxial half, scabrous; palea narrow, 2-keeled, 2.5–3.5 mm, keels rough, hyaline. Anthers 1–1.5 mm, pale orange to reddish. Caryopsis 0.5–1 mm, fusiform, brown. Illustration. Parodi (1949: 446). Chromosome number. 2n 5 14 (Hubbard, 1984). Distribution, habitat, and phenology. Deschampsia setacea is known from bogs and wet places in western Europe, from Scandinavia to the Iberian Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:53 157 Cust # 2008115 158 Annals of the Missouri Botanical Garden Peninsula. In South America, it is found in central Chile in similar habitats; it has also been cited for southern Chile (Parodi, 1949), but no material from this region has been found. It grows between 900 and 3500 m elevation. Flowering occurs in January and February. Discussion. Deschampsia venustula and D. patula are similar in having glumes that are violaceous along the keels and hyaline or light green toward the margins. Deschampsia venustula differs from D. patula in its longer, slightly more contracted panicles (up to 14 3 4.5 cm vs. wider, less contracted panicles up to 10 3 6 cm in D. patula) and in the awns, which exceed the glumes in D. venustula and are included in D. patula. Discussion. Deschampsia setacea is similar to Avenella flexuosa in the bristlelike leaves, but the two differ in the longer and pointed ligules of D. setacea (vs. obtuse in Avenella) and in the more purple-colored spikelets of A. flexuosa. Specimens examined. CHILE. Coquimbo Region: Ovalle, Quebrada Larga, C. Jiles 4143 (CONC); Illapel, Rı́o Ojotas, near La Vega Redonda, J. Morrison & R. Wagenknecht 17424 (BAA, SI). Santiago Metropolitan Region: Las Condes, cordillera al oriente de Santiago, Mina Disputada, G. Looser 1111 (BAA, CONC); A. Garaventa 544 (BAA); Cordillera de las Arañas, Jan. 1861, G. Land s.n. (SGO 045880). 15. Deschampsia venustula Parodi, Darwiniana 8: 450. 1949. TYPE: Chile. Cordillera de Santiago, Valle Largo, Feb. 1892, F. Philippi s.n. (holotype, SGO not seen; isotypes, BAA ex SGO!, US 556484 ex SGO not seen, photo!). Perennial, caespitose, densely tufted, erect, culms 5–27 cm tall, 1-noded, sheaths glabrous, with membranous margins. Leaf blades setaceous, 1– 4.5 cm 3 0.5–1 mm, nerves on both sides usually glabrous, sometimes with prickles on the adaxial side; ligules acute, 3–7 mm, scarious, margins of the ligule prolonged into the upper part of the sheaths. Panicles slightly contracted, 3–14 3 1.5–4.5 cm, with 5 to 7 verticils, branches scabrous, purplish. Spikelets 2-flowered, erect; callus and rachilla pilose, callus trichomes short, barely reaching the middle of the lemma; lower glume narrowly lanceolate to lanceolate, 3–4 mm, 1-nerved, upper glume lanceolate, 3.5– 4.5 mm, obscurely 3-nerved, both glumes normally scabrous and purplish only along the keels, pale gold toward the margins, scarious; lemma 4-toothed, lateral teeth larger than the central tooth, 2.5–3 mm, 4(5)nerved, nerves glabrous; awn bent and twisted, 4– 5 mm, scabrous, inserted at or near the base, exceeding the glumes; palea 2-keeled, hyaline, rarely membranous, keels scabrous. Anthers 0.5–1 mm, brownish red. Caryopsis 0.5–1 mm, fusiform. Illustration. Nicora (1978: 234). Distribution, habitat, and phenology. Deschampsia venustula occurs in the Andes of Argentina and Chile, from 32uS to ca. 38uS, in humid and open places at altitudes between 200 and 4300 m. Flowering occurs between January and March. Specimens examined. ARGENTINA. Mendoza: Dpto. San Carlos, Cordillera del Portillo de La Llareta (entre El Paso del Portillo y la Laguna del Diamante, F. Kurtz 10991 (CORD [sheets A, B]); Laguna Diamante, G. Covas 1047 (BAB, SI, US); próximo a la laguna, O. Boelcke 4122 (BAA, BAB); J. Hueck 18191 (SI). Neuquén: Dpto. Chos Malal, cajón del Arroyo del Cruce, 36u439S, 70u239W, O. Boelcke et al. 11272 (BAB). CHILE. Santiago Metropolitan Region: Santiago, Estero del Plomo, La Disputada, 33u079S, 70u219W, M. T. K. Arroyo 83-1340 (CONC). CONCLUSIONS Deschampsia in South America comprises 15 species, some of which show a high degree of morphological similarity, requiring additional studies to clarify their true relationships. In all cases, the species can be recognized, but the existence of intergrading forms suggests a close relationship, which, if confirmed, might result in changes in specific status (i.e., some taxa reduced to subspecies or varieties). Groups of species with difficult circumscription are: D. kingii and D. laxa; D. antarctica, D. parvula, D. patula, and D. venustula; and D. cespitosa, D. cordillerarum, and D. mendocina. Two monotypic genera formerly included in Deschampsia are also found in the region: Avenella flexuosa and Vahlodea atropurpurea. The placement of Deschampsia is conflictive because molecular-based analyses (Soreng & Davis, 2000; Quintanar et al., 2007; Davis & Soreng, 2007; Soreng et al., 2007) did not support its placement inside the ‘‘true’’ Aveneae, but in a closely related clade called ‘‘former Aveneae lineages related to the traditional Poeae’’ (Quintanar et al., 2007: 1563). The fact that molecular results often collide with traditional morphological classifications is well known (Brummit, 2006; Middleton, 2006). The differing data from morphological and molecular studies suggest two possible placements for Deschampsia: (1) including Deschampsia in the traditional Aveneae, based on morphological data, or (2) including them in the Poeae, based on molecular data. A classificatory system that reconciles morphological and molecular data is still lacking, but at this point it seems that the inclusion of Deschampsia and allies in the Aveneae as Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:53 158 Cust # 2008115 Volume 97, Number 2 2010 traditionally defined is no longer possible. The more appropriate placement—at the moment—is tribe Poeae, subtribe Airinae Fr., as advocated by Soreng et al. (2007). Concerning the origin of Deschampsia, Raven and Axelrod (1974) and Stebbins (1975) suggested a northern origin for all the pooid tribes (including the Aveneae s.l. and former Aveneae). Kawano (1963, 1966) hypothesized on the basis of cytological data that the most common species (D. cespitosa) might have covered the area of Pleistocene glaciation in the Northern Hemisphere almost completely, and that the postglacial migration of surviving populations into the glaciated lands may have occurred several times. Long-distance dispersal events have probably been involved in the migration of ancestors to the south, where a center of secondary differentiation (Tateoka, 1962; Hartley, 1973) developed as a result of the existence of temperate climatic conditions similar to those of the north. The limited molecular data available (Chiapella, 2007) suggest that although Deschampsia is monophyletic, some differences exist between northern and southern populations of the widespread D. cespitosa. Based on biogeographical evidence, Hartley (1973) found the Aveneae s.l. to be more diverse and abundant in the Northern Hemisphere, but, based on the present distribution of the species, Deschampsia has flourished in the Southern Hemisphere, where it has developed an important center of diversity in South America. EXCLUDED OR UNCERTAIN TAXA Deschampsia brasiliensis (Louis-Marie) Valencia, Revista Argent. Agron. 8: 128. 1941. Basionym: Trisetum brasiliense Louis-Marie, Rhodora 30: 242. 1928 [1929]. TYPE: Brazil. Rio de Janeiro, Alto de Itatiaia, dense tufts in peaty soils among rocks, above timberline, 17 Jan. 1925, 2200–2400 m, A. Chase 8304 (holotype, US 1257235 not seen, photo!; isotypes, BAA fragm. ex US!, MO 924156 not seen, photo!). The fragment conserved at BAA presents isolated spikelets, mostly 1-flowered, with the florets with stout awns that are not typical of Deschampsia. It is not possible to clearly observe the lemmas with the 4-toothed apex, which is one of the few extremely constant characters in the genus. The species was excluded from Trisetum (Finot et al., 2005: 535), so its generic placement remains unclear. Deschampsia conferta (Pilg.) Valencia, Revista Argent. Agron. 8: 127. 1941. Basionym: Trisetum confertum Pilg., Bot. Jahrb. Syst. 25(5): 714. 1898. Peyritschia conferta (Pilg.) Finot, Contr. U.S. Natl. Herb. 48: 478. 2003. TYPE: Ecuador. ‘‘Loma de Canaballa y alrededores, Provincia Imbabura,’’ 2100–2300 m, 1 Feb. Chiapella & Zuloaga Revision of Deschampsia, Avenella, and Vahlodea 159 1871, A. Stübel 152 (holotype, B not seen, photo!; isotype, US 81771 not seen, photo!). This taxon is excluded from Deschampsia because of the lemmas, which are awnless or awned with bilobed apex, in opposition to Deschampsia, which has awned lemmas with 4-toothed apex. The plant has inflorescences in narrow, contracted panicles as described for Peyritschia E. Fourn. by Finot et al. (2004). The specimen cited by Jorgensen and Ulloa Ulloa (1994), E. Asplund 6976 (S!), collected in Pichincha, Ecuador, agrees well with Peyritschia. Deschampsia latifolia Phil., Linnaea 29: 91. 1858, nom. illeg. hom. TYPE: Chile. Andes de Linares, s.d., Germain 197 (holotype, SGO PHIL 197 not seen, SGO photo!; isotypes, BAA fragm. ex SGO PHIL 197!, US 556494 fragm. ex SGO PHIL 197 not seen, photo!). The fragment conserved in BAA consists of a few spikelets with glumes ca. 7 mm long, and the lemmas have no awn or teeth. The leaf blades are ca. 10 mm wide, while leaf blades are usually no more than 5 mm wide in South American Deschampsia. Deschampsia micrantha Phil., Anales Univ. Chile 94: 24. 1896. TYPE: Chile. Coquimbo, Entrada de Tilito, 7 Feb. 1883, F. Philippi s.n. (holotype, SGO 37497 not seen, photo!; isotypes, BAA!, SGO 62687 not seen, photo!, SGO 63067 not seen, photo!, US 865603 fragm. ex SGO not seen, photo!). The specimen conserved in BAA is a complete plant, ca. 42 cm high, with a basal compact tuft of conduplicate, almost filiform leaf blades no longer than 5 cm, an open panicle ca. 9 3 5 cm, and 1-flowered spikelets. The lemmas are similar to Deschampsia, but are erose at the apex rather than 4-toothed. The ligular zone presents triangle-shaped thickenings similar to the lateral pulvini described by Rúgolo (1986) for Deyeuxia Clarion ex P. Beauv. Literature Cited Albers, F. 1972a. Cytotaxonomie und B-Chromosomen bei Deschampsia caespitosa (L.) P.B. und verwandten Arten. Beitr. Biol. Pflanzen 48: 1–62. ———. 1972b. Cytosystematische Untersuchungen in der Subtribus Deschampsiineae Holub (Tribus Aveneae Nees). II. Die Gattungen Vahlodea Fr. und Avenella Koch. Ber. Deutsch. Bot. Ges. 85: 279–285. ———. 1978. Karyologische und genomatische Veränderungen innerhalb der Gräser-Subtriben Aristaveninae und Airinae. Ber. Deutsch. Bot. Ges. 91: 693–697. ———. 1980a. Systematik, variation und Entwicklungstendenzen der Subtriben Aristaveninae und Airinae (Gramineae–Aveneae). Phyton (Horn) 20: 95–116. ———. 1980b. Vergleichende Karyologie der GräserSubtriben Aristaveninae und Airinae (Poaceae–Aveneae). Pl. Syst. Evol. 136: 137–167. ———. 1980c. Karyogeobotanik der Gräser-Subtriben Aristaveninae und Airinae (Aveneae). Flora 169: 150–167. Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:54 159 Cust # 2008115 160 Annals of the Missouri Botanical Garden ——— & F. Butzin. 1977. Taxonomie und Nomenklatur der Subtriben Aristaveninae und Airinae (Gramineae–Aveneae). Willdenowia 8: 81–84. Arancio, G., M. Muñoz & F. A. Squeo. 2001. Descripción de algunas especies con problemas de conservación en la IV Región de Coquimbo, Chile. Pp. 63–103 in F. A. Squeo, G. Arancio & J. R. Gutiérrez (editors), Libro Rojo de la Flora Nativa y de los Sitios Prioritarios para su Conservación: Región de Coquimbo. Ediciones Universidad de La Serena, La Serena, Chile. Arroyo, M. T. K., J. J. Armesto & C. Villagran. 1981. Plant phenological patterns in the High Andean Cordillera of central Chile. J. Ecol. 69: 205–223. ———, M. Marticorena, O. Matthei, M. Muñoz & P. Pliscoff. 2002. Analysis of the contribution and efficiency of the Santuario de la Naturaleza Yerba Loca, 33uS in protecting the regional vascular plant flora (Metropolitan and Fifth regions of Chile). Revista Chil. Hist. Nat. 75: 767–792. Beetle, A. A. 1987. Las Gramı́neas de México, Vol. 2. Secretarı́a de Agricultura y Recursos Hidráulicos. Comisión Técnico Consultiva de Coeficientes de Agostadero (COTECOCA), México D.F. Bor, N. L. 1970. Gramineae. In K. L. Reichinger (editor), Flora des Iranischen Hochlandes und der Umrahmenden Gebirge, Persien, Afghanistan, Teile von West-Pakistan, Nord Iraq, Azerbaidjan, Turkmenistan. Akademische Druck- und Verlagsanstalt, Graz. Brummit, R. K. 2006. Am I a bony fish? Taxon 55: 268–269. Buschmann, A. 1949. Zwei für Europa neue adventive Deschampsia-Arten. Phyton (Horn) 1: 190–193. Campbell, C. S., J. A. Quinn, G. P. Cheplick & T. J. Bell. 1983. Cleistogamy in grasses. Ann. Rev. Ecol. Syst. 14: 411–441. Castlebury, L. A., L. M. Carris & K. Vánky. 2005. Phylogenetic analysis of Tilletia and allied genera in order Tilletiales (Ustilaginomycetes; Exobasidiomycetidae) based on large subunit nuclear rDNA sequences. Mycologia 97: 888–900. Cheeseman, T. F. 1906. Manual of the New Zealand Flora. John Mackay, Wellington. Chiapella, J. 2000. The Deschampsia cespitosa complex in central and northern Europe: A morphological analysis. Bot. J. Linn. Soc. 134: 495–512. ———. 2007. A molecular phylogenetic study of Deschampsia (Poaceae: Aveneae) inferred from nuclear ITS and plastid trnL sequence data: Support for the recognition of Avenella and Vahlodea. Taxon 55: 55–64. ———. 2009. Neotypification of Aira setacea Hudson (Poaceae). Watsonia 27: 239–242. ——— & N. S. Probatova. 2003. The Deschampsia cespitosa complex (Poaceae: Aveneae) with special reference to Russia. Bot. J. Linn. Soc. 142: 213–228. Clarke, G. C. S. 1980. Deschampsia. Pp. 225–227 in T. G. Tutin, V. H. Heywood, N. A. Burges, D. M. Moore, D. H. Valentine, S. M. Walters & D. A. Webb (editors), Flora Europaea, Vol. 5. Cambridge University Press, Cambridge. Clayton, W. D. 1970. Gramineae (part 1). In E. MilneRedhead & R. M. Polhill (editors), Flora of Tropical East Africa. Crown Agents for Oversea Governments and Administrations, London. ——— & S. A. Renvoize. 1986. Genera Graminum. Kew Bull. Addit. Ser. XIII. Conert, H. J. 1987. Deschampsia. Pp. 302–317 in Gustav Hegi (editor), Illustrierte Flora von Mitteleuropa, Band 1, Teil 3. Parey Buchverlag, Berlin. Corte, A. 1961. Fertilidad de las semillas en fanerógamas que crecen en Cabo Primavera (costa de Danco), Penı́nsula Antárctica. Contr. Inst. Antarc. Argent. 65: 1–16. Cracraft, J. 1983. Species concept and speciation analysis. Curr. Ornithol. 1: 159–187. Cronquist, A., A. H. Holmgren, N. H. Holmgren, J. L. Reveal & P. K. Holmgren. 1977. Intermountain Flora, Vol. 6. Columbia University Press, New York. Davis, J. I. & R. J. Soreng. 2007. A preliminary phylogenetic analysis of the grass subfamily Pooideae (Poaceae), with attention to structural features of the plastid and nuclear genomes, including an intron loss in GBSSI. Aliso 23: 335–348. Davidse, G. 1994. Deschampsia. Pp. 888–999 in G. Davidse, M. Sousa S. & A. O. Chater (editors), Flora Mesoamericana, Vol. 6: Alismataceae a Cyperaceae. Universidad Nacional Autónoma de México, Mexico D.F.; Missouri Botanical Garden, St. Louis; The Natural History Museum, London. Desvaux, E. 1854. Gramineas. In C. Gay (editor), Flora de Chile 6: 233–469; 342, tab. 79, fig. 1. Du Rietz, G. 1930. The fundamental units of biological taxonomy. Svensk. Bot. Tidskr. 24: 333–428. Dusén, P. 1900. Die gefässpflanzen der Magellansländer: Beitrage zur Flora der Ostküste von Patagonien. Svenska Exped. Magellansländerna 3: 77–264. Finot, V. L., P. M. Peterson, R. J. Soreng & F. O. Zuloaga. 2004. A revision of Trisetum, Peyritschia, and Sphenopholis (Poaceae: Pooideae: Aveninae) in Mexico and Central America. Ann. Missouri Bot. Gard. 91: 1–30. ———, ———, F. O. Zuloaga, R. J. Soreng & O. Matthei. 2005. A revision of Trisetum (Poaceae: Pooideae Aveninae) in South America. Ann. Missouri Bot. Gard. 92: 533–568. Frey, L. 1999. Avenella—A genus of Aveneae (Poaceae) worthy of recognition. Fragm. Florist. Geobot. 7(Suppl.): 27–32. Garcia-Suarez, R., C. Alonso-Blanco, M. C. FernandezCarvajal, J. A. Fernandez-Prieto, A. Roca & R. Giraldez. 1997. Diversity and systematics of Deschampsia sensu lato (Poaceae), inferred from karyotypes, protein electrophoresis, total genomic DNA hybridization and chloroplast DNA analysis. Pl. Syst. Evol. 205: 99–110. Haraldsen, K. B., M. Ødegaard & I. Nordal. 1991. Variation in the amphi-Atlantic plant Vahlodea atropurpurea (Poaceae). J. Biogeogr. 18: 311–320. Hartley, W. 1973. Studies on the origin, evolution, and distribution of the Gramineae. V. The subfamily Festucoideae. Aust. J. Bot. 21: 201–234. Hauman, L. 1918. La végétation des Hautes Cordillères de Mendoza. Anales Soc. Ci. Argent. 86: 121–188. Hickman, J. C. 1993. The Jepson Manual: Higher Plants of California. University of California Press, Berkeley. Hitchcock, C. L., A. Cronquist, M. Ownbey & J. W. Thompson. 1969. Vascular Plants of the Pacific Northwest, Part 1: Vascular Cryptogams, Gymnosperms and Monocotyledons. University of Washington Press, Seattle. Holderegger, R., I. S. Stehlik, R. I. L. Smith & R. J. Abbott. 2003. Populations of antarctic hairgrass (Deschampsia antactica) show low genetic diversity. Arctic Alpine Res. 35: 229–235. Holmgren, A. H. & N. H. Holmgren. 1977. Poaceae. In A. Cronquist, A. H. Holmgren, N. H. Holmgren, J. L. Reveal & P. K. Holmgren (editors), Intermountain Flora, Vol. 6. Columbia University Press, New York. Holmgren, P. K., N. H. Holmgren & L. C. Barnett. 1990. Index Herbariorum, ed 8. New York Botanical Garden, Bronx. Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:54 160 Cust # 2008115 Volume 97, Number 2 2010 Chiapella & Zuloaga Revision of Deschampsia, Avenella, and Vahlodea 161 Hooker, J. D. 1847. The Botany of the Antarctic Voyage of H.M. Discovery ships Erebus and Terror in the Years 1839–1843. J. Cramer (reprinted 1963), Weinheim. Hubbard, C. E. 1984. Grasses: A Guide to Their Structure, Identification, Uses, and Distribution in the British Isles, 3rd ed. Revised by J. C. E. Hubbard. Penguin Books, Harmondsworth. Hultén, E. 1941. Flora of Alaska and Yukon. C. W. Gleerup, Lund. ———. 1968. Flora of Alaska and Neighboring Territories. Stanford University Press, Stanford. Jørgensen, P. M. & C. Ulloa Ulloa. 1994. Seed Plants of the High Andes of Ecuador: A Checklist. Aalborg University Reports No. 34. Kawano, S. 1963. Cytogeography and evolution of the Deschampsia cespitosa complex. Canad. J. Bot. 41: 719–742. ———. 1966. Biosystematic studies of the Deschampsia cespitosa complex with special reference to the karyology of Icelandic populations. Bot. Mag. (Tokyo) 79: 293–307. Linnaeus, C. 1753. Species Plantarum. Holmiae, Stockholm. Macloskie, G. 1904. Reports of the Princeton University Expeditions to Patagonia 1896–1899. 8: 139–338. The University, Princeton. McNeill, J., F. R. Barrie, H. M. Burdet, V. Demoulin, D. L. Hawksworth, K. Marhold, D. H. Nicolson, J. Prado, P. C. Silva, J. E. Skog, J. H. Wiersema & Turland (editors). 2006, International Code of Botanical Nomenclature (Vienna Code). Regnum Veg. 146. Middleton, D. J. 2006. Nomenclatural notes on N. J. Asian Apocynaceae, subfamilies Rauvolfioideae and Apocynoideae. Taxon 55: 502–506. Moberg, R. & Ö. Nilsson. 1991. Typification of Nordic vascular plants. 1. Names published by G. Wahlenberg. Nordic J. Bot. 11: 287–299. Nicora, E. 1978. Poaceae. Pp. 225–238 in M. N. Correa (editor), Flora Patagónica, Vol. 8(3). Instituto Nacional de Tecnologı́a Agropecuaria, Buenos Aires. Nixon, K. C. & Q. D. Wheeler. 1990. An amplification of the phylogenetic species concept. Cladistics 6: 211–233. Ortiz-Troncoso, O. R. 1976. A 16th century Hispanic harbour in the Strait of Magellan, South America. Int. J. Naut. Archaeol. 5: 176–179. Palisot de Beauvois, A. M. F. J. 1812. Essai d’une nouvelle Agrostographie; ou nouveau genres des Graminées; avec figures représentant les caracters de tous les genres. Paris (VI). Parodi, L. R. 1949. Las gramı́neas sudamericanas del género Deschampsia. Darwiniana 8: 415–475. Philippi, R. A. 1858. Plantarum novarum Chilensium V. Linnaea 29: 48–95. ———. 1864. Plantarum Novarum Chilensiam Centuriae, inclusis quibusdam Mendocinis et Patagonicis. Linnaea 33: 1–308. ———. 1873. Memorias cientı́ficas i literarias. Botánica. Anales Univ. Chile 43: 479–583. ———. 1896. Plantas Nuevas Chilenas. Anales Univ. Chile 94: 5–34. Porter, D. M. 1986. Charles Darwin’s vascular plant specimens from the voyage of HMS Beagle. Bot. J. Linn. Soc. 93: 1–172. Quicke, D. L. J. 1993. Principles and Techniques of Contemporary Taxonomy. Blackie Academic & Professional, London. Quintanar, A., S. Castroviejo & P. Catalán. 2007. Phylogeny of the tribe Aveneae (Pooideae, Poaceae) inferred from plastid trnT-F and nuclear ITS sequences. Amer. J. Bot. 94: 1554–1569. Raven, P. H. & D. I. Axelrod. 1974. Angiosperm biogeography and past continental movements. Ann. Missouri Bot. Gard. 61: 539–673. Rúgolo de Agrasar, Z. 1986. Las especies extrapatagónicas del género Deyeuxia (Gramineae): II. Morfologı́a de la lı́gula de Deyeuxia chrysantha Presl. Parodiana 4: 73–95. Rzedowski, J. & G. Rzedowski. 1990. Flora Fanerogámica del Valle de Mexico 2. Publicaciones del Instituto de Ecologia, Michoacán. Soreng, R. J. & J. I. Davis. 2000. Phylogenetic structure in Poaceae subfamily Pooideae as inferred from molecular and morphological characters: Misclassification versus reticulation. Pp. 61–74 in S. W. L. Jacobs & J. Everett (editors), Grasses: Systematics and Evolution. Commonwealth Scientific and Industrial Research Organisation, Collingwood, Victoria. ———, ——— & M. A. Voionmaa. 2007. A phylogenetic analysis of Poaceae tribe Poeae sensu lato based on morphological characters and sequence data from three plastid-encoded genes: Evidence for reticulation, and a new classification for the tribe. Kew Bull. 62: 425–454. Stebbins, G. L. 1975. The role of polyploid complexes in the evolution of North American grasslands. Taxon 24: 91–106. Tateoka, T. 1962. Notes on some grasses. XIV. Enumeration of genera of Festuceae. Bot. Mag. (Tokyo) 75: 336–343. Tzvelev, N. N. 1976. Grasses of the Soviet Union. Nauka Publishing House, Leningrad. Valencia, J. 1941. Especies crı́ticas de Trisetum que deben pasar al género Deschampsia. Revista Argent. Agron. 8: 122–130. van de Vouw, M., P. van Dijk & A. H. L. Huiskes. 2007. Regional genetic diversity patterns in Antarctic hairgrass (Deschampsia antarctica Desv.). J. Biogeogr. 35: 365–376. Vigo i Bonada, J. 1983. Flora de la Vall de Ribes. Acta Bot. Barcinon. 35: 1–793. APPENDIX 1. Examined material of Deschampsia. Each specimen is cited by the last name of the first collector when there is more than one collector. Species number is indicated between parentheses and corresponds to the List of Species below. LIST OF SPECIES 1. 2. 3. 4a. 4b. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. Deschampsia Deschampsia Deschampsia Deschampsia Deschampsia pulchra Deschampsia Deschampsia Deschampsia Deschampsia Deschampsia Deschampsia Deschampsia Deschampsia Deschampsia Deschampsia Deschampsia airiformis (Steud.) Benth. & Hook. f. antarctica E. Desv. berteroana (Kunth) Trin. cespitosa (L.) P. Beauv. var. cespitosa cespitosa (Nees & Meyen) Nicora var. cordillerarum Hauman danthonioides (Trin.) Munro elongata (Hook.) Munro kingii (Hook. f.) E. Desv. laxa Phil. looseriana Parodi mendocina Parodi parvula (Hook. f.) E. Desv. patula (Phil.) Pilg. ex Skottsb. setacea (Huds.) Hack. venustula Parodi Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:54 161 Cust # 2008115 162 Annals of the Missouri Botanical Garden Abrams 2833 (7); Alboff 1027 (2), s.n. (12); Allen 38 (7); Ambrosetti 206 (2), 1202 (3), 7713 (3); Arneberg 9632 (8); Arroyo 83-1340 (15), 85-155 (13), 87-0007 (12), s.n. (2). Barnier 232 (10), Barrientos 11 (10), 220 (7), 511 (3), 530 (3), 976 (3), 1637 (10), 1655 (3), 3308 (3), 5867 (7), 5871 (7), 9913 (3); Beetle 1731 (6); Behn 12493 (2); Biurrun 5280 (4a); Blankinship 599 (7); Boelcke 3201 (1), 4122 (15), 4172 (4b), 4486 (2), 6051 (7), 9765 (5), 10213 (4a), 12358 (13), 12446 (2), 13830 (4b), 14056 (1), 14330 (4a); Boelcke et al. 11272 (15); Bonarelli 78 (9), 96 (2); Bossieu s.n. (2); Bravo 23 (10); Burkart 6211 (4a), 9507 (2), 9529 (7); Burmeister s.n. (13), s.n. (8). Cabrera 6169 (4b); Calder 29623 (6), 35255 (7); Carette 22 (4a); Castellanos 12831 (12), 7572 (2), 12846 (8), s.n. (4a); Collantes 2107 (13); Correa 1923 (2), 3082 (2), 5598 (7); Corte 1 (2); Covas 1047 (15). Dauber 168, 196 (2); De Giorgio 452 (3); De La Rüe s.n. (2); Dimitri 2843 (4a), 4455 (4a), 8243 (2); Durán 3507 (6); Dusén 405 (2), 548 (8), 549 (7). Elmer 1664 (7); Eyerdam 24115 (2), 24132 (8). Faggi s.n. (4a), s.n. (7); Feruglio s.n. (4a); Figueroa 3094 (4a). Garaventa 544 (14), 1667 (3), 2838 (3); Gehrling 109 (4a); Goodall 4700 (8); Grandjot 15a (2), 3486a (4a); Greene 3058 (2); Grondona 4480 (2), 7226 (12); Guiñazú 183 (2); Gunckel 18244 (3), 20285 (3), 25067 (3), 26730 (3); Gusinde 474 (8). Haene 1979 (4a); Hauman s.n. (4a); Heller 3289 (7); Hitchcock 1417 (7), 1419 (6); Hoover 6789 (6); Hueck 18191 (15); Hunziker, A. T. 8218 (8), 8225 (2), 10112 (2), 10121 (2), 10147 (2), 10150 (2), 10152 (2), 10153 (2), 10190 (2), 10196 (2), 10200 (2), 10205 (2); Hunziker, J. H. 6709 (8), 6763 (2). Illı́n 167 (7), 224 (7), 226 (4a); Iter Patagonicum 126 (12). Jara s.n. (7); Jiles 3623 (5), 4143 (14), 6073 (3); Johnson 586 (9); Johnston 5962 (4a), 6096 (4a); Jones 2157 (6). Kalela 2021 (8), 2022 (7); Kildale 6100 (7); Koslowsky 142 (8), 224 (2); Kunkel 631 (1), 2004 (3); Kurtz 32 (8), 9495a (4b), 9667 (4a), 9727 (4a), 10991 (15). Laffuel 1838 (10), 1903 (2); Land s.n. (14); Laurı́a s.n. (2); León 2400 (13); Levi 2674 (10); Longton 601 (2); Looser 1111 (14), 1384 (3), 3438 (10), 3718 (10); Luti 1435 (8), 1485 (2). Maldonado 129 (7); Marticorena 143 (6), 216 (6), 228 (6), 387 (10); Martı́nez 2 (2); Martı́nez 52495 (3); Martı́nez Crovetto 3028 (7); Matthei 1167 (3); McVaugh 10323 (7); Mexı́a 7978 (8); Montero 2366 (10); Moore 276 (2), 781 (2), 1349 (2), 1654 (12), 1688 (8), 1758 (12), 2800 (12); Moore Jr. 3126 (7); Morrison 17424 (14). Navas 8123 (10); Nicora 753-17 (7), 3635 (7), 3768 (4a), 9340 (1), 10231 (2), 26417 (2). Parodi 9550 (8), 11425 (7), 15358 (1), s.n. (8); Pastore 72 (8); Paz Martı́nez 7 (4a); Pennington 187 (2), 267 (8), 392 (2); Pérez Moreau s.n. (4a), s.n. (9), s.n.(4a); Pfister 7383 (7), 11938 (2), 13102 (6); Philippi s.n. (3); Pisano 5089 (8), 5123 (13), 5474 (9), 7577 (12), 8191 (7); Popovici s.n. (2); Pöppig s.n. (3); Pringle 4743 (7), 13244 (7); Procter 16 (2). Reynoso-Muller 3864 (4a); Ricardi 375 (7); Ritter 2199 (4a); Roig 77 (13); Ru 9695 (4a); Ruiz Leal 7843 (4a), 15025 (2); Ryves 96/081 (3). Santana Michel 3411 (7); Schlegel 2590 (4a); Seijo 1735 (4a); Siple 336 (2); Skottsberg 63 (2), s.n. (8); Sladen H639/1 (2); Smith 13371 (4a), 15713 (4a); Soriano 1511 (4a), 2245 (2), 2554 (2), 3083 (2), 3196½ (2), 4474 (2), 4626 (1), 5368 (8), 5415 (2), 5802 (1); Spegazzini 20701 (12). TBPA 1912 (2), 2139 (2), 2916 (12), 3284 (8), 3647 (12), 5103 (9), 5472 (8); Tiehm 14258 (6); Tomé s.n. (10); Tortorelli s.n. (8); Turquets s.n. (2). Ulibarri 1067 (2). Vallerini 322 (7), 3900 (13); Valls 2394 (4a). Werdermann 582 (4a); Werth s.n. (2). Zöllner 9623 (12), 46212 (5). Annals of the Missouri Botanical Garden mobt-97-02-01.3d 20/5/10 12:56:58 162 Cust # 2008115

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A REVISION OF DESCHAMPSIA, AVENELLA, AND VAHLODEA (POACEAE, POEAE, AIRINAE) IN SOUTH AMERICA

A REVISION OF DESCHAMPSIA, AVENELLA, AND VAHLODEA (POACEAE, POEAE, AIRINAE) IN SOUTH AMERICA

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