Contributions from the United States National Herbarium Volume 56: 1-184 A Revision of Fes tuca (Poaceae: Loliinae) in South American Paramos Daniel Stancik and Paul M. Peterson Smithsonian Institution Scholarly Press A Revision of Fes tuca (Poaceae: Loliinae) in South American Paramos Daniel Stancik and Paul M. Peterson Smithsonian Institution Scholarly Press WASHINGTON, D.C. 2007 ABSTRACT Stancik, Daniel and Paul M. Peterson. A revision of Festuca (Poaceae: Loliinae) in South American paramos. Smithsonian Contributions from the United States National Herbarium, Volume 56, 184 pages (including 97 figures). A taxonomic revision of Festuca L. for the South American paramos of Brazil, Colombia, Ecuador, northern Peru, and Venezuela is given. Fifty-six species and six subspecies of Festuca are recognized in the study area. Fifty-three species are native to the flora region and three are introduced and cultivated. Keys for determining the species, a classification, synonymies, descriptions, leaf anatomical descriptions, observations, distribution and habitat, specimens examined, illustrations, and leaf surface micrographs are provided. One species: F. turimiquirensis Stancik & P.M. Peterson, and one subspecies: F. tolucensis subsp. culata Stancik & P.M. Peterson, are newly described. The following three names were lectotypified: Bromus caldasii Kunth [=Festuca caldasii (Kunth) Kunth], Festuca fratercula Rupr. ex E. Fourn. (=Festuca amplissima Rupr. subsp. amplissima), and Festuca ovina subvar. jamesonii St.-Yves (=Festuca glumosa Hack, ex E.B. Alexeev). KEY WORDS: Anatomy, Brazil, Classification, Colombia, Ecuador, Festuca, Loliinae, Poaceae, Taxonomy, Venezuela. RESUMEN Stancik, Daniel and Paul M. Peterson. A revision of Festuca (Poaceae: Loliinae) in South American paramos. Smithsonian Contributions from the United States National Herbarium, xxx paginas (incluyendo 97 figuras). Se presenta una revision taxonomica del genero Festuca (Poaceae) para los paramos del Sudamerica cubriendo la region de Brazil, Colombia, Ecuador, norte del Peru y Venezuela. Se reconocen 56 especies y seis subespecies. Cincuenta y tres especies son nativas de la region, tres son introducidas y cultivadas. En el trabajo se presenta una clave para la determination de las especies, una clasificacion del genero en la region estudiada, sinonimos, descripciones, description de la anatomia foliar, distribution y ecologia de las especies, lista de los especimenes estudiados, ilustraciones y microfotografias de la epidermis foliar. Se describe una nueva especie: F. turimiquirensis Stancik & P.M. Peterson, y una subespecie F. tolucensis subsp. culata Stancik & P.M. Peterson. Tambien se designan lectotipos para tres nombres: Bromus caldasii Kunth [=Festuca caldasii (Kunth) Kunth], Festuca fratercula Rupr. ex E. Fourn. (=Festuca amplissima Rupr. subsp. amplissima) y Festuca ovina subvar. jamesonii St.-Yves (=Festuca glumosa Hack, ex E.B. Alexeev). PALABRAS CLAVES: Anatomia, Brasil, Clasificacion, Colombia, Ecuador, Festuca, Loliinae, Poaceae, Taxonomia, Venezuela. Published by Smithsonian Institution Scholarly Press P.O. Box 37012 MRC 957 Washington, D.C. 20013-7012 www.scholarlypress.si.edu The periodical Contributions from the U.S. National Herbarium was first published in 1890 by the United States Department of Agriculture. From July 1,1902, forward it was published as a Bulletin of the United States National Museum. The series was discontinued after volume 38(1974) and revived with volume 39 (2000) as a venue for publishing taxonomic papers, checklists, floras, and monographs produced by staff and associates at the U.S. National Herbarium. It is externally peer reviewed and published at irregular intervals. Subscriptions and other correspondence should be addressed to Smithsonian Institution Scholarly Press. The present issue is available in print for free while supplies last; PDF files are available at http://www.sil.si.edu/smithsoniancontributions. Contributions from the United States National Herbarium (ISSN 0097-1618) Cover design by Alice R. Tangerini; front, habit of Festuca fimbriata Nees; back, habit of Festuca fragilis (Luces) B. Briceno. Illustrated by Anna Skoumalova. POSTMASTER: Send address changes to Smithsonian Institution Scholarly Press, P.O. Box 37012, MRC 957, Washington, D.C. 20013-7012. Library of Congress Cataloging-in-Publication Data Stancik, Daniel. A revision of festuca (Poaceae: Loliinae) in South American paramos / Daniel Stancik and Paul M. Peterson, p. cm. • (Contributions from the United States National Herbarium ; 56) Includes bibliographical references and index. Summary in Spanish. 1. Fescue•South America•Classification. 2. Fescue•South America•Identification. I. Peterson, Paul M. II. Title. QK495.G74S728 2007 584'.9•dc22 2007026985 S9 The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48•1992. CONTENTS INTRODUCTION TAXONOMY AND DIVERSITY 5 5 6 BlOGEOGRAPHY 6 HISTORICAL OVERVIEW MATERIAL AND METHODS ACKNOWLEDGEMENTS REFERENCES TAXONOMIC TREATMENT KEY TO THE SPECIES OF FESTUCA IN BRAZIL, COLOMBIA, ECUADOR, AND VENEZUELA SURVEY OF THE SPECIES AND SUBGENERIC TAXA OF FESTUCA IN SOUTH AMERICAN PARAMOS DESCRIPTIONS OF SPECIES OF FESTUCA FROM SOUTH AMERICAN PARAMOS EXCLUDED NAME SCANNING ELECTRON MICROGRAPHS OF LEAF EPIDERMAL SURFACES APPENDICES INDEX TO FIGURES ALPHABETICAL INDEX OF ACCEPTED SPECIES AND SUBSPECIES ALPHABETICAL INDEX TO COLLECTORS INDEX TO SCIENTIFIC NAMES 7 8 8 11 11 16 17 146 147 173 173 175 176 183 A Revision of Festuca (Poaceae: Loliinae) in South American Paramos Daniel Stancik1 and Paul M. Peterson2 INTRODUCTION Composed of more than 400 species, Festuca is one of the largest genera in the Poaceae (Clayton & Renvoize 1986; Watson & Dallwitz 1992). The genus has cosmopolitan distribution and it is an important component of grass ecosystems of the temperate zone as well as the alpine grasslands of the tropical zone. Although the diversity of the genus Festuca is centered in the Holarctic zone of Eurasia and North America, with about 140 species (Darbyshire et al. 2003), the South American Andes represent a significant center of Festuca diversity. HISTORICAL OVERVIEW.•In South America, taxonomic studies of Festuca have been concentrated in the southern part of the continent, i.e. Argentina, Bolivia, Chile, and Peru (Phillipi 1859-60, 1865; Pilger 1898, 1906; Hitchcock 1927; Saint-Yves 1927; Parodi 1935,1953; Macbride 1936; Infantes 1952; Foster 1966; Tiirpe 1969; Vareschi 1970; Correa 1978; Matthei 1982; Tovar 1993; Brako & Zarucchi 1993; Zuloaga et al. 1994; Renvoize 1998; Soreng et al. 2003). Taxonomic information on Festuca in the paramo zone of South America (Venezuela, Colombia, Ecuador, N Peru, SE Brazil) has been quite sporadic, although botanical collections have increased in the last 50 years. The oldest specimens of Festuca from Venezuela were collected during 1906-1911 by A. Jahn, H. Pittier. The first list of Venezuelan grasses was published in 1937 (Anonymous 1937) and only two species of Festuca (F. livida and F tolucensis) were listed. In 1942, Luces distinguished between the Mesoamerican and the Venezuelan populations of F. livida and described a new Venezuelan species, Ffragilis (Luces) Briceno. Two species {Ffragilis and F tolucensis) were mentioned in the first Venezuelan paramo flora by Vareschi (1970). Later, Alexeev (1986) in his work on Festuca for Venezuela, Colombia, and Ecuador mentioned only one species from Venezuela (F tolucensis). The recently published study of Briceno and Murillo (1994) treats Festuca in the state Merida (Venezuela) and lists five native species (F. pinetorum Swallen, F elviae Briceno, F tolucensis, Ffragilis, F coromotensis Briceno). Stancik and Peterson (2003) indicated that Fpinetorum was misidentified and therefore does not occur in Venezuela. In addition, Davidse et al. (1994) mentioned the species F. amplissima Rupr.; the only known Venezuelan specimen (Tillett & Honig 746-765, MO) was collected in Serrania de Perija on the ColombianVenezuelan boundary. The earliest collections of Festuca from Ecuador and Colombia were made by A. von Humboldt and A. J. A. Bonpland (Expedition to Ecuador in 1802-1803; see Humboldt et al. 1816 and Jorgensen et al. 1999) and J.C. Mutis and F.J. Caldas (Real Expedicion Botanica 1783-1808; see Pinto-Escobar 1976,1985; and Blanco & del Valle 1991). Seven species are present in Humboldt's collection, five of which were later described by C.S. Kunth as new (F. andicola, F caldasii, F. dasyantha, F procera, F. quadridentata). The Festuca specimens collected by T. Haenke (Expedicion Malaspina 1789-1794) are of even earlier date, but none of them are from Ecuador or Colombia. The uncertain origin of the type specimen of Festuca presliana Hitchc. (= Bromus depauperatus J. Presl) is discussed in Stancik & Peterson (2003). Hitchcock (1927), in his overview of grasses of Ecuador, Peru, and Bolivia, mentioned 17 species. The Ecuadorian species included^ dasyantha, ' Department ofBotany, Charles University of Prague, Bendskd 2, 128 01 Prague 2, Czech Republic, dan_stancik@yahoo.com 2 Department of Botany MRC-166, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA. peterson@si.edu Festuca in South American Paramos F. ulochaeta, F. eminens, F procera, F sublimes, F. andicola, F. dichoclada, F. dolichophylla, F. breviaristata. Festuca eminens is also cited for Colombia. Hitchcock's publication represents the first overview of grasses for this region but does not reflect the entire diversity, since more than 60 species are known from the region today. Additionally, the accepted "broad species" concept results in the inclusion of species (such as F ulochaeta, F dichoclada, F. dolichophylla), that do not appear in Ecuador. The detailed work of Saint-Yves (1927) has little practical use for Festuca taxonomists in South America, as the work suffers from insufficient specimens studied and appears to be focused on minute details rather than estimating species variability and pointing out differences. The taxa were recognized in a confused system of varieties and forms. Later, descriptions of new species were published only occasionally (Swallen 1948) and as checklists of local floras and herbaria collections (Pinto-Escobar 1966,1985; Acosta-Solis 1969; Burger 1981; Cleef 1981). There treatments mechanically accepted the conclusions of previous authors (principally of Hitchcock) and confusion surrounding species delimitations have continued (e.g., the citation of F. dolichophylla for Colombia). New insight into the taxonomy of Festuca in northern South America was brought by Alexeev (1984,1986), particularly in terms of lectotypification of previously described species and description of new taxa at sub-generic and specific levels. Alexeev (1986) cited 19 native species from Ecuador and Colombia. The work was based on the examination and revision of a limited number of herbarium specimens without the advantage of field work and it could not reflect the entire diversity of the genus in South America. Alexeev's conclusions were reflected in the checklists of local floras published later (Jorgensen & UlloaUlloa 1994; Jorgensen & Leon Yanez 1999; Luteyn 1999; Rangel 2000). Interest in the study of the Brazilian paramo is recent (Safford 1999a, b). TAXONOMY AND DIVERSITY.•Fifty-three native and three introduced species are recognized in the study area. Among them: five species (F dinirica, F. guramacalana, F hatico, F. turimiquirensis and F. venezuelana) and two subspecies (F tolucensis subsp. culata and F tolucensis subsp. perijae) are recognized as new for Venezuela; twelve species (F boyacensis, F. chita, F. chitagana, F. cocuyana, F hatico, F monguensis, F nereidaensis, F pilarfranceii, F. toca, F sanctae-martae, F sumapana, and F. woodii) and one subspecies (F. amplissima subsp. magdalenaensis) as new for Colombia; six species (F carchiense, F holubii, F imbaburensis, F laegaardii, F oroana, and F. soukupii) and three subspecies (F chimborazensis subsp. micacochensis, F parciflora subsp. loxana, and F vaginalis subsp. cayambae) as new to Ecuador; and, one species (F. renvoizii) as new to Peruvian jalca (i.e., humid alpine grass ecosystem of the northern Peru). The species found in South America are attributed to the six subgenera and seven sections of the genus Festuca. About 80% of South American Festuca species belong to the large cosmopolitan subg. Festuca. Remarkably, three subgenera {Festuca subg. Helleria, subg. Mallopetalon, and subg. Erosiflorae) and three sections {Festuca sect. Ruprechtia, sect. Glabricarpae, and sect. Cataphyllophorae) are endemic or largely endemic to South America. The distribution of species diversity is not homogeneous in South America. The Andes between central Peru and northern Argentina has the highest species concentration (almost 50% of all South American taxa). In the northern Andes, there are two areas with high species diversity, central Ecuador (Pichincha with 14 species, and Chimborazo with 13 species) and northern Colombia (Boyaca with 14 species). High levels of local endemicity were observed in regions with the highest species diversity, especially in Boyaca. Similarity of species composition was also analyzed to elucidate the relationships among the South American regions. As expected, strong relationships exist between the Festuca species of Ecuador and Colombia, and the Festuca species from northern South America ("paramo group") are clearly distinct from the those of central and southern South America. BIOGEOGRAPHY.•The known range of several Festuca species is expanded contributing to a better understanding of the biogeography of the genus in South America. Festuca amplissima was found in northern Colombia (Sierra Nevada de Santa Marta) and on the Colombian-Venezuelan boundary (Serrania de Perija). This species represents the biogeographical Festuca in South American Paramos connection between Mesoamerican and South American flora. The species was originally known from Costa Rica, Guatemala, Mexico, and Panama. Festuca fragilis was found in Nevado del Cocuy (Dept. Boyaca, Colombia) and was previously reported only from the state of Merida in Venezuela, ca. 250 km from the Colombian locality. Festuca ulochaeta was found in the Cordillera Oriental of the Colombian Andes as well as the northern Andes and Cordillera de la Costa of Venezuela, and in Costa Rica. Formerly, this species was known only from the region of Brazilian Shield of southeastern Brazil and northeastern Argentina. This information might be important in determining the origins and phylogeny of other South American species of Festuca sect. Subulatae. The first collections of the genus Festuca in Colombian Cordillera Occidental are reported: F. asplundii (Farallones de Cali and Macizo de Tamana), F. sodiroana (Paramo Frontino), and F andicola (Paramo Frontino). Festuca caldasii, F glumosa, and F sodiroana, which were formerly regarded as endemic to Ecuador, were confirmed in Colombia, showing the biogeographical relationship between Colombian Cordillera Central and Ecuadorian Andes. Similarity, Festuca asplundii, which was originally considered endemic to Colombia and Ecuador, was confirmed in the Andes of northern Peru. Festuca dinirica and F. guaramacalana are the first records of the genus reported from the mountains of National Park Dinira and from the massif of National Park Guaramacal in Venezuela, respectively, and represent the most easterly Andean records of Festuca. Important extra-Andean findings of Festuca in Venezuela were made in Cordillera de la Costa en Venezuela. The localities Colonia Tovar and Serrania de Turimiquire represent the most easterly, mountain forests that include Festuca in northern South America. These localities are located 250 km and 600 km east of the nearest Andean locality, respectively. The following species that were formerly mentioned in the floras of Venezuela, Colombia, Ecuador, or Peru were not confirmed: F andicola (excluded from Peru), F casapaltensis Ball (excluded from Ecuador), F. dasyantha Kunth (excluded from Colombia), F. dichoclada Pilg. (excluded from Ecuador), F dolichophylla J. Presl (excluded from Colombia and Ecuador), F ortophylla Pilg. (excluded from Ecuador), Fperuviana Infantes (excluded from Ecuador), Fpinetorum (excluded from Venezuela), F. quadridentata Kunth (excluded from Colombia), F rigescens (J. Presl) Kunth (excluded from Ecuador), F. vaginalis (Benth.) Laegaard (excluded from Colombia), and F. ulochaeta Nees ex Steud. (excluded from Ecuador). MATERIAL AND METHODS The work presented here summarizes the results of taxonomic and biogeographic research on the genus Festuca in paramos of South America that was carried out between 1998 and 2004. Extensive field and herbarium studies were performed to assess the diversity of the genus in this geographic area of South America. The area studied included Brazil, Colombia, Ecuador, northern Peru, and Venezuela. An exhaustive bibliographic search for literature on the taxonomy of Festuca was performed, with special focus on the genus in Mesoamerica and South America. The protologues, local floras, and monographs were consulted to document all reports of Fectuca from this area. More than 1,500 specimens of Festuca from the study area and 1,000 specimens from other Mesoamerican and South American countries were studied. Specimens are located at AAU, B, BA, BAA, BOG, BOL, CAUP, C, CAR, COL, CUVC, CTES, F, FAUC, FMB, HUA, HERZ, JAUM, JBGP, K, LOJA, LP, LPB, MA, MEDEL, MER, MERC, MERF, MY, NY, QCA, QCNE, QAP, QPLS, P, PORT, PR, PRC, PSO, R, RB, TOLI, TULV, U, UIS, UPTC, US, UTMC, VALLE, VEN, and W (abbreviations according to Holmgren et al. 1998 onwards). The type specimens of all recognized species and nearly all types of their synonyms were examined. The taxonomic analysis is based on extensive morphological and anatomical comparisons. Twenty-six morphological characters were selected as useful for the descriptions and differentiation among the species. The selected characters were: height of plants, formation of shoot innovations, number and position of the nodes, presence/absence of auricles, length and diameter of the leaf blades, Festuca in South American Paramos shape and length of ligule, color and structure of veins, vestiture of culms, length of panicle, length of spikelets (measured total length of spikelet) and number of florets, length, shape and vestiture of glumes, lemma and palea, length of anthers, and length of hilum. After preliminary observations (Stancik 1999), five anatomical characters of the leaf cross sections (number of vascular bundles, number of ribs, distribution of the sclerenchyma, relative density, and length of trichomes) were also included in the study. Anatomical sections were prepared from dry herbarium specimens after hydratation in a soap solution and observed under microscope at 100x magnification. Ecuatoriano Ciencias Naturales, Contribution 71, Quito. Alexeev, E.B. 1984. The new taxa of the genus Festuca from Colombia and Ecuador. Bot. Zhurn. 69(11): 1543-1551 (in Russian). Alexeev, E.B. 1986. Festuca L. (Poaceae) in Venezuela, Colombia et Ecuador. Novosti Sist. Vyss. Rast. 23: 5-23 (in Russian). Anonymous. 1937. Lista Provisional de las Gramineae senaladas en Venezuela hasta 1936. Ministerio de Agricultura. Boletin Tecnico 1, Caracas. ACKNOWLEDGEMENTS We would like to thank the Faculty of Science, Charles University of Prague, for institutional support during realization of this research. We also thank the grant agency of Czech Republic (projects GACR 206/03/0197, GACR 206/06/P293), Ministry of Education of Czech Republic (project MSM 0021620828), and the Smithsonian Institution for financial support. Our thanks belong also to Anna Skoumalova for providing plant illustrations. We wish to thank the Laboratory of Electron Microscopy (LEM) of the Institute of Microbiology, Academy of Sciences of the Czech Republic (Dr. O. Benda, O. Kofronova, majority of photos on Fig. 72-97), LEM of Aarhus University (A. Sloth - Fig. 76E, 77A,B, 85C,D, 89C,D, 94A,B, 97C), LEM of Royal Botanic Gardens Kew (Dr. H. Banks - Fig.74A-D, 93E, F) for assistance during preparation of micro photos. This study could not have been completed without the kind assistance of the curators and researchers at the following herbaria: AAU, B, BA, BAA, BOG, BOLV, CAUP, C, CAR, COL, CONC, CUVC, CTES, FAUC, FMB, GB, HUA, HERZ, JAUM, JBGP, K, L, LOJA, LP, LPB, MA, MEDEL, MER, MERC, MERF, MO, MY, NY, QCA, QCNE, QAP, QPLS, P, PORT, PR, PRC, PSO, R, RB, SGO, TOLI, TULV, U, UIS, UPTC, US, UTMC, VALLE, VEN, W Finally, the Smithsonina Institution Scholarly Press is acknowledged for preparing the final version for printing. Blanco and del Valle, A. 1991. Herbarium Mutisianum. Fontqueria 32: 1-173. Brako, L. and J.L. Zarucchi.1993. Catalogue of the Flowering Plants and Gymnosperms of Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 45: 1-1286. Briceno, B. and G. Morillo. 1994. El genero Festuca L. (Poaceae) en los paramos de Merida. Ernstia. 4(3-4): 73-88. Clayton, WD. and S.A. Renvoize. 1986. Genera Graminum. Grasses of the world. Kew Bull., Addit. Ser. 13: 1-389. Cleef, A.M. 1981. 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Journal of Biogeography 26: 713-737. Saint-Yves, A. 1927. Contribution a l'etude des Festuca (subg. Eu-Festuca) de l'Amerique du Sud. Candollea3: 151-315. Salamanca, V.S. 1991. The vegetation of the paramo and its dynamics in the volcanic massif RuizTolima. (Cordillera Central Colombiana). Ph.D. Thesis, University of Amsterdam, The Netherlands. 122 pp. Sanchez, M.R. and J.O. Rangel. 1990. Estudios ecologicos en la cordillera Oriental Colombiana V. Analisis fitosociologico de la vegetacion de los depositos turbosos paramunos de los alrededores de Bogota. Caldasia 16(77): 155-192. Sodiro, PL. 1889. Gramineas ecuatorianas de la provincia de Quito. Anales de la Universidad de Quito 3(25): 474-484. Soreng, R.J, P.M. Peterson, G. Davidse, E.J. Judziewicz, F.O. Zuloaga, T.S. Filgueiras, and O. Morrone. 2003. Catalogue of New World grasses (Poaceae) IV. Subfamily Pooideae. Contr. U.S. Natl. Herb. 48: 1-730. Stancik, D. 1999. La problematica del uso de las secciones transverzales de hojas en el estudio taxonomico de Festuca L. (Poaceae) en Colombia. In: Rangel Ch., J.O. et al., eds. Libro de resumenes. 1st Congreso Colombiano de Botanica. ICN UN. Bogota. 160 pp. Stancik, D. 2004. New taxa of Festuca (Poaceae) from Ecuador. Folia Geobot. 39: 97-110. Stancik, D. and P.M. Peterson. 2003. Festuca dentiflora (Poaceae: Loliinae: sect. Glabricarpae), a new species from Peru and taxonomic status of Fpresliana. Sida20(3): 1015-1022. Sturm, H. and J.O. Rangel Ch. 1985. Estudios ecologicos en los paramos andinos: Una vision preliminar integrada. Biblioteca J.J. Triana 9: 167-224. ICN Univ. Nacional de Colombia. Swallen, J. R. 1948. New greasses from Honduras, Colombia, Venezuela, Ecuador, Bolivia, and Brazil. Contr. U.S. Natl. Herb. 29(6): 251-276. Tovar, O. 1972. Revision de las especies peruanas del genero Festuca, Gramineae. Mem. Mus. Hist. Nat. "Javier Prado" 16: 1-93. Tovar, O. 1993. Las gramineas (Poaceae) del Peru. Ruizia 13: 1-480. Turpe, A. M. 1969. Las especies argentinas de Festuca. Darwiniana. 15(1-2): 189-283. Vareschi, V 1970. Flora de los paramos de Venezuela. Edic. Univ. de los Andes. Merida, 404-407. Vargas, R.O. and S. Zuluaga. 1985. La vegetacion del paramo de Monserrate. In: Sturm, H. and J.O. Rangel Ch., eds. Estudios ecologicos en los paramos andinos: Una vision preliminar integrada. Biblioteca J.J. Triana 9: 167-224. Verveij, PA. 1995. Spatial and temporal modeling of vegetation patterns: Burning and grazing Festuca in South American Paramos in the paramo of Los Nevados National Park, Colombia. Ph.D. Thesis, University of Amsterdam, The Netherlands. Watson, L. and M.J. Dallwitz. 1992. The Grass Genera of the World. Cambridge University Press, Cambridge. 1036 pp. 11 Zuloaga, F. O., E.G. Nicora, Z.E. Riigolo de Agrasar, O. Morrone, J. Pensiero, and A.M. Cialdella. 1994. Catalogo de la familia Poaceae en la Republica Argentina. Monogr. Syst. Bot. Missouri Bot. Gard. 47: 1-178. TAXONOMIC TREATMENT L., Sp. PI. 1: 73. 1753. TYPE: F. ovina L., Sp. PI. 1: 73-74. 1753. (lectotype: designated byNash,Ill.Fl.US.Canad.,ed.2,1:269.1913; also Jarvis et al., Watsonia 16: 300. 1987). FESTUCA Perennial herbs, densely or loosely caespitose (forming often large tussocks) or rhizomatous, commonly monoecious, exceptionally dioecious (not in South America). Culms unbranched 0.1-2 m high, below panicle glabrous or finely scabrous, with 1 or more (2-6) glabrous nodes. Innovations extravaginal, intravaginal or mixed. Sheaths with free or partially united margins, non-auriculate (except in Festuca subg. Schedonorus); basal sheaths occasionally thickened into a bulb; ligules membranous; normally less than 1 mm, rarely to 10 mm long; leaf blades flat but often conduplicate or setaceous and then basal, without cross nerves, 0.2-15 mm wide, basal cataphyls rarely present. Inflorescences an open or contracted panicle. Spikelets usually with 2 or several bisexual or unisexual florets and an apical rudiment, chasmogamous, some species cleistogamous or viviparous i.e., vegetative proliferation of the spikelets, laterally compressed, disarticulating above the glumes and below the florets; glumes lanceolate to ovate, pointed, carinate or non-carinate, shorter than adjacent lemma; lower glume shorter, 1 (rarely 3)-nerved; upper glume 3 (rarely 5)-nerved; lemmas commonly membranous (rarely coriaceous), rounded (exceptionally carinate), 5-nerved, entire (rarely bidentate), acute, shortly mucronate or awned; awn continuous with midnerve, terminal, rarely subterminal, up to 15 mm long; paleas membranous, two-carinate, two-dentate, as long as the lemma or a little shorter, scabrous at keels, awnless; lodicules two, small, about 1 mm long, two-dentate, hyaline.; stamens 3, anthers 0.4-6 mm long; ovaries glabrous, rarely hairy at apex, styles terminal without an apical appendage; stigmas 2, white. Fruit a caryopsis, free or with lemma adherent, with a linear or sometimes oblong hilum. Leaf blade anatomy.•Cross sections with five or more vascular bundles, with or without adaxial ribs over the vascular bundles; bulliform cells present or absent; sclerenchyma under abaxial epidermis continuous or discontinuous, adaxial sclerenchyma fascicles present or absent; sclerenchyma sometimes extending to the vascular bundles abaxially and also sometimes adaxially forming girders; hairs on adaxial epidermis dense or scattered, 20-200 urn long, prickles on abaxial epidermis present or absent. KEY TO THE SPECIES OF FESTUCA IN BRAZIL, COLOMBIA, ECUADOR, AND VENEZUELA la. Plants rhizomatous; culms solitary 30-100 cm tall or loosely caespitose herbs with culms 80-250 cm tall; plants often in mountain forests 2 lb. Plants densely caespitose, without rhizomes (only several species exceptionally forming short rhizomes inside the tussocks); culms short 8-50(-60) cm tall or culms tall 60-170 cm tall; plants often in mountain grasslands 24 2a. Cataphylls (basal shoots coriaceous or with coriaceous-membranous scales) present 2b. Cataphylls absent 3a. Panicle branches densely hairy; ligules l-1.5(-2) mm long 3b. Panicle branches glabrous or scabrous; ligules less than 1 mm long 3 6 27. F. laegaardii 4 12 Festuca in South American Paramos 4a. Spikelets with 4-6 florets; lower glumes 2-3 mm long; upper glumes 3-4 mm long; lemmas 4-5 mm long, awned; paleas as long as adjacent lemma 30. F. toca 4b. Spikelets with 2•4 florets; lower glumes more than 3 mm long; upper glumes more than 4 mm long; lemmas more than 5 mm long, short-awned; paleas shorter than adjacent lemma 5 5a. Panicles nutant and nodding, 9-12 cm wide; lower glumes 4-5 mm long; upper glumes 5.5-6.5 mm long; lemmas 6-7 mm long; anthers 1-1.6 mm long 23. F. chitagana 5b. Panicles erect, 0.5-1 cm wide; lower glumes 3-3.6 mm long; upper glumes 4.5-5 mm long; lemmas 6-6.5 mm long; anthers 3.5 mm long 26. F. hatico 6a. Auricles large, more than 1 mm long, falcate 6b. Auricles absent or small, less than 1 mm long and inconspicuous, not falcate 7a. Margins of auricles hairy; upper glumes 4.5-6.5 mm long 7b. Margins of auricles without hairs; upper glumes 3.5-4 mm long 7 8 12. F. arundinacea 13. F. pratensis 8a. Apex of lodicules hairy 8b. Apex of lodicules glabrous 18. F. fimbriata 9 9a. Lower glumes 0.8-2.5 mm long 9b. Lower glumes longer than 2.5 mm long 10 15 10a. Lemma awns more than 4 mm long 10b. Lemma awns less than 4 mm long 11a. Ligules 3-3.5 mm long; lemma awns 9-12 mm long lib. Ligules 1-2 mm long; lemma awns 5-7 mm long 12a. Spikelets 10-13 mm long; lemmas 7-8.5 mm long; anthers 3-3.5 mm long 12b. Spikelets 7-10 mm long; lemmas 4.5-6.5 mm long; anthers 0.8-1.5 mm long 11 12 4. F. flacca 7. F. tovariensis 10. F. reclinata 13 13a. Leaf blades flat; panicles nutant; lemmas awnless 6. F. sodiroana 13b. Leaf blades conduplicate or involute, sometimes flat; panicles erect; lemmas with awns 0.51.5 mm 14 14a. Culms solitary with long rhizomes; glumes and lemmas purple; panicles 0.5-1 cm wide 19. F. andicola 14b. Culms loosely tufted; glumes and lemmas green; panicles 1-2 cm wide 3. F. elviae 15a. Ligules 2.5-13 mm long 15b. Ligules 0.5-1.5(-2) mm long 16 19 16a. Ligules 7-13 mm long; spikelets 11-14 mm long; lemmas awnless 16b. Ligules 2.5-5 mm long; spikelets 14-17 mm long; lemma awns 1-8 mm long 17 18 17a. Lower glumes (3.5-)4•5 mm long; upper glumes 5-6.5 mm long; margins of lemmas erose, toothed 15. F. quadridentata 17b. Lower glumes 5-7.5 mm long; upper glumes 7-8 mm long; margins of lemma entire 14. F. dichoclada 18a. Upper glumes 4.5-6.5 mm long; lemmas 10-14 mm long; anthers 3.5^4.5 mm long 9. F. caldasii Festuca in South American Paramos 13 18b. Upper glumes 7-9 mm long; lemmas 10-11 mm long; anthers 3-3.5 mm long 16. F. venezuelana 19a. Lower glumes 2.5-3.5(-4) mm long 19b. Lower glumes 4.5-6 mm long 20 21 20a. Leaf blades 3-5 mm wide; upper glumes 4.5-6 mm long; lemmas awnless or with awns 0.5 mm long, straight 2. F. coromotensis 20b. Leaf blades 5-11 mm wide; upper glumes 3.5^4.5 mm long; lemma awns 7-15 mm long, flexuous 8. F. ulochaeta 21a. Upper glumes 5-5.5 mm long; lemmas 6.5-7 mm long; awns 2.5-3.5 mm long 21b. Upper glumes 6-8 mm long; lemmas 8-10 mm long; lemma awns 0-1 mm long 20. F. rubra 22 22a. Leaf blades conduplicate or involute, 0.6-0.7 mm in diameter; ligules 1-1.5 mm long; lemmas 9.5-10 mm long 5. F. guaramaca 22b. Leaf blades flat, 3-12 mm wide; ligules 0.5-1 mm long; lemmas 8-9 mm long 23 23a. Caespitose plants with rhizomes, 120-180 cm tall; lemma awnless; anthers 3 mm 1. F. amplissima 23b. Culms solitary, rhizomatous, 70-90 cm tall; lemma short-awned, the awns 0.5-1 mm long; anthers 3.5^4 mm long 11. F. woodii 24a. Cataphylls (basal shoots coriaceous or with coriaceous-membranous scales) present 24b. Cataphylls absent 25a. Lemmas with a dense covering of long hairs 25b. Lemmas glabrous or scabrous 26a. Leaf blades 0.8-1.6 mm in diameter; abaxial schlerenchyma continuous 26b. Leaf blades 0.4-0.7 mm in diameter; abaxial schlerenchyma discontinuous 25 30 25. F. dasyantha 26 27 29 27a. Cataphylls 2-5 cm long; ligules 0.1-0.3 mm long; lemmas 4.5-6 mm long 28. F. pilar-franceii 27b. Cataphylls 0.5-1.5 cm long; ligules more than 0.3 mm long; lemmas 6-8 mm long 28 28a. Culms 70-80 cm tall; panicles contracted, 1-6 cm wide; spikelets 9-10 mm long 22. F. azucarica 28b. Culms 80-170 cm tall; panicles lax, 15-25 cm wide; spikelets 11-15 mm long 29. F. procera 29a. Lower glumes 5.5-6.5 mm long; upper glumes 7-8.5 mm long; anthers 0.8-1.1 mm long 35. F. chita 29b. Lower glumes 3^4.5 mm long; upper glumes 4.5-6.5 mm long; anthers 2-3.5 mm long 24. F. colombiana 30a. Plants growing in large tussocks, culms (30-)50-100 cm tall and 0.3-1 m in diameter near base 31 30b. Plants growing in small tussocks, culms 15-60(-80) cm tall and 0.05-0.2 m in diameter near base 43 31a. Leaf blades scabrous 31b. Leaf blades glabrous 32 36 14 Festuca in South American Paramos 32a. Leaf blades 0.5-0.7 mm in diameter; ligules predominantly acute 32b. Leaf blades 0.8-1.5 mm in diameter; ligules truncate 33 35 33a. Some spikelets transformed into vegetative shoots (viviparous plants), sexual organs in such spikelets absent 52. F. subulifolia 33b. Spikelets perfect, well developed, no spikelets vegetatively prolifering 34 34a. Spikelets 8-14 mm long; lower glumes 6-8.5 mm long; upper glumes 6.5-9.5 mm long; lemmas 6-10 mm long 54. F. tolucensis 34b. Spikelets 6.5-9 mm long; lower glumes 3.5-5.5 mm long; upper glumes 4-6 mm long; lemmas 5-6.5(-8) mm long 52. F. subulifolia 35a. Spikelets perfect, not vegetatively prolifering, 8-10 mm long, with 3^1 florets; lemmas 6.5-8.5 mm long, with awns 0.7-1 mm long 37. F. densipaniculata 3 5b. All spikelets with vegetative proliferation; lemmas transformed into vegetative shoots, sexual organs absent 29. F. asplundii 3 6a. Leaf blades flat or conduplicate, 2-5 mm in diameter; spikelets 12-15 mm long 42. F. glyceriantha 36b. Leaf blades involute, less than 2 mm in diameter; spikelets shorter than 12 mm long 37 37a. Anthers 1•1.5 mm long 37b. Anthers 2-3 mm long 38 39 38a. Culms about 50 cm tall; leaf blades 0.6-0.8 mm in diameter; spikelets with 3-4 florets; lower glumes 4 mm long; upper glumes 5 mm long 46. F. monguensis 38b. Culms 80-100 cm tall; leaf blades 1.2-1.5 mm in diameter; spikelets with 5 florets; lower glumes 6-6.5 mm long; upper glumes 7-8 mm long 47. F. nereidaensis 39a. Leaf blades l.l-1.8(-4) mm in diameter; lower glumes 2.4-2.6 mm long; upper glumes 3.63.9 mm long 48. F. oroana 39b. Leaf blades 0.4-1 mm in diameter; lower glumes 3-7 mm long; upper glumes 4-8 mm long.... 40 40a. Ligules longer than 1 mm; panicles open 40b. Ligules shorter than 1 mm; panicles contracted 41 42 41a. Ligules (1.5•)2•3.5 mm long; lower glumes 3-4 mm long; upper glumes 4-5.5 mm long 55. F. turimiquensis 41b. Ligules (0.5-)l-2(-2.5) mm long; lower glumes (3.5-)4-6(-6.5) mm long; upper glumes (5.5-) 6-7.5(-8) mm long 36. F. cleeflana 42a. Leaf blades 0.4-0.6 mm in diameter; ligules 0.5-1 mm long; spikelets with 3-4 florets; lemmatal awns 1.5•4 mm long 40. F. dinirica 42b. Leaf blades 0.7-0.1 mm in diameter; ligules 0.1-0.3 mm long; spikelets with 5-7(-8) florets; lemmatal awns 0-1 mm 38. F. cundinamarcae 43a. Spikelets about 3 cm long, with 5-7 florets; lemmas 17-20 mm long 43b. Spikelets less than 2 cm long, lemmas less than 15 mm long 17. F. fragilis 44 44a. Lower glumes 1.2-2 mm long; upper glumes 2-2.7 mm long; lemmas 4.5-5.5 mm long 21. F. soukupii Festuca in South American Paramos 15 44b. Lower glumes longer than 2 mm; upper glumes longer than 2mm; lemmas longer than 15 mm 45 45a. Leaf blades glabrous 45b. Leaf blades scabrous 46 52 46a. Lemma with awns longer than 1 mm 46b. Lemma awnless or awns shorter than 1 mm 47 48 47a. Culms 8-15(-30) cm tall; lower glumes 2.5-3.5 mm long; upper glumes 3.5-5 mm long; lemma with awns 1-2 mm 34. F. chimborazensis 47b. Culms 15-55 cm tall; lower glumes 4-4.5 mm long; upper glumes 5.5-6.5 mm long; lemma with awns 4-6 mm 44. F. huamachucensis 48a. Leaf blades less than 0.8 mm in diameter; lower glumes 2.5-4.5 mm long; upper glumes 4-6.5 mm long; lemmas shorter than 7.5 mm 49 48b. Leaf blades (0.5-)0.8-3.5 mm in diameter; lower glumes 6.5-10 mm long; upper glumes 6.510.5 mm long; lemmas 7.5-10 mm 50 49a. Ligules 0.5-0.7 mm long; upper glumes 4-4.2 mm long; lemmas 5.5-6(-7) mm long 45. F. imbaburensis 49b. Ligules 1.2-1.5 mm long; upper glumes 5-5.5 mm long; lemmas 7-7.5 mm long 50. F. renvoizii 50a. Leaf blades involute, 0.5-0.8 mm in diameter; panicles about 0.5 cm wide; spikelets 8-9.5 mm long; lemmas 7.5-8.5 mm long 56. F. vaginalis 50b. Leaf blades conduplicate, up to 3.5 mm wide; panicles 0.8-2 cm wide; spikelets 10-13 mm; lemmas 8.5-10 mm 51 51 a. Culms 20-50 cm tall; leaf blades 0.8-2 mm in diameter; ligules 0.6-1 mm long; panicles 0.8-1.5 cm wide; lower glumes 7.5-8 mm long 37. F. cocuyana 51b. Culms 60-80 cm tall; leaf blades 2-3.5 mm wide; ligules 1-1.6 mm long; panicles 1.5-2 cm wide; lower glumes 8.5-9.5 mm long 51. F. sanctae-martae 52a. Ligules 0.3-0.5 mm long;callus and rachilla glabrous 52b. Ligules 0.5-1.2 mm long; callus and rachilla scabrous or sparsely hairy 53 54 53a. Culms 15-20 cm tall; spikelets 9.5-11 mm long; lower glumes 4-5 mm long; upper glumes 5.5-6 mm long; paleas as long as adjacent lemma; anthers 0.6-0.8 mm long 53. F. sumapana 53b. Culms 30-60 cm tall; spikelets 8-9.5 mm long; lower glumes 3-3.7 mm long; upper glumes 4-5 mm long; paleas shorter than adjacent lemma; anthers 0.9-1.2 mm long 32. F. boyacensis 54a. Leaf blades 0.4-0.7 mm in diameter with 5 vascular bundles; paleas as long as adjacent lemma... 55 54b. Leaf blades 0.8-1.4 mm in diameter, with 7 vascular bundles; paleas shorter than adjacent lemma 56 55a. Leaf blades glaucous; spikelets 8.5-9 mm long; lower glumes 3-3.5 mm long; upper glumes 4-5 mm long 49. F. parciflora 55b. Leaf blades green; spikelets 9-10.5 mm long; lower glumes 3.5-4.5 mm long; upper glumes 5-6.5 mm long 33. F. carchiense 16 Festuca in South American Paramos 56a. Culms 50-60 cm tall; ligules 0.6-0.8 mm long; lower glumes 4 mm long; upper glumes 5.5 mm long; anthers 1.3 mm long 43. F. holubii 56b. Culms 15-55 cm tall; ligules 0.8-1.2 mm long; lower glumes 4.5-6 mm long; upper glumes 6-6.5 mm long; anthers 0.8-0.9 mm long 41. F. glumosa SURVEY OF THE SPECIES AND SUBGENERIC TAXA OF FESTUCA IN SOUTH AMERICAN PARAMOS 1. Subg. Drymanthele V.I. Krecz. & Bobrov, Fl. URSS 2: 532. 1934. TYPE: F. drymeja Mert. & W.D.J. Koch 1.1. Sect. Ruprechtia E.B. Alexeev, Novosti Sist. Vyss. Rast. 17: 42. 1980. TYPE: F. amplissima Rupr. 1. Festuca amplissima 2. Subg. Subulatae (Tzvelev) E.B. Alexeev, Bjull. Moskovsk. Obsc. Isp. Prir, Otd. Biol. 82(3): 96. 1977. TYPE: F subulata Trin. 2.1. Sect. Subulatae Tzvelev, Bot. Zhurn. 56(9): 1253. 1971. TYPE: F subulata Trin. 2. Festuca coromotensis 3. Festuca elviae 4. Festuca flacca 5. Festuca guaramaca 6. Festuca sodiroana 1. Festuca tovariensis 8. Festuca ulochaeta 2.2. Sect. Glabricarpae E.B. Alexeev, Bot. Zhurn. 67(9): 1291. 1982. TYPE: F. breviglumis Swallen 9. Festuca caldasii 10. Festuca reclinata 11 Festuca woodii 3. Subg. Schedonorus (P. Beauv.) Peterm., Deutschl. Fl. 643. 1849. TYPE: F arundinaceae Schreb. 3.1. Sect. Schedonorus (P. Beauv.) Koch, Syn. Fl. Germ. Helv. 810. 1837. TYPE: F arundinaceae Schreb. 12. Festuca arundinacea 13. Festucapratensis 4. Subg. Erosiflorae E.B. Alexeev, Novosti Sist. Vyssh. Rast. 23: 11. 1986. TYPE: F quadridentata Kunth 14. Festuca dichoclada 15. Festuca quadridentata 16. Festuca venezuelana 5. Subg. Helleria E.B. Alexeev, Novosti Syst. Vyssh. Rast. 17: 42. 1980. TYPE: F livida (Kunth) Willd. ex Spreng. 17. Festucafragilis 6. Subg. Mallopetalon (Doll) E.B. Alexeev, Bot. Zhurn. 69(3): 346. 1984. TYPE: Ffimbriata Nees. 18. Festucafimbriata 7. Subg. Festuca 7.1. Sect. Aulaxyper Dumort, Observ. Gramin. Belg.: 104. 1824. TYPE: F. rubra L. 19. Festuca andicola 20. Festuca rubra 21. Festuca soukupii 7.2. Sect. Cataphyllophorae E.B. Alexeev, Bot. Zhurn. 69(11): 1546. 1984. TYPE: F procera Kunth 22. Festuca azucarica 23. Festuca chitagana 24. Festuca colombiana 25. Festuca dasyantha 26. Festuca hatico Festuca in South American Paramos 17 27. Festuca laegaardii 28. Festucapilar-franceii 29. Festuca procera 30. Festuca toca 7.3. Sect. Festuca 31. Festuca asplundii 32. Festuca boyacensis 33. Festuca carchiense 34. Festuca chimborazensis 35. Festuca chita 36. Festuca cleefiana 37. Festuca cocuyana 38. Festuca cundinamarcae 39. Festuca densipaniculata 40. Festuca dinirica 41. Festuca glumosa 42. Festuca glyceriantha 43. Festuca holubii 44. Festuca huamachucensis 45. Festuca imbaburensis 46. Festuca monguensis 47. Festuca nereidaensis 48. Festuca oroana 49. Festuca parciflora 50. Festuca renvoizei 51. Festuca sanctae-martae 52. Festuca subulifolia 53. Festuca sumapana 54. Festuca tolucensis 55. Festuca turimiquirensis 56. Festuca vaginalis DESCRIPTIONS OF SPECIES OF FESTUCA FROM SOUTH AMERICAN PARAMOS 1. Festuca amplissima Rupr. subsp. amplissima, Bull. Acad. Roy. Sci. Bruxelles 9(2): 236.1842. (Figs. 2,72A-D). TYPE: Mexico. Veracruz, Pic d'Orizaba, 10,000 ft, Jun-Oct 1840,//. Galeotti 57(5(5 (lectotype: BR; isotype: W!). Festuca fratercula Rupr. ex E. Fourn. Mexic. PI. 2:124. 1886. TYPE: Mexico. Veracruz, Cordillera, Pico de Orizaba, 12,000 ft, Jun-Oct 1840, H. Galeotti 5778 (lectotype: LE!, designated here). UniolaeffusaE. Fourn.,Mexic. PL 2:122.1886. TYPE: Mexico. San Nicolas, E. Bourgeau 1032 (holotype: P; isotype: US-865711 fragm. ex P!). Uniola muelleri E. Fourn., Mexic. PI. 2: 122.1886. TYPE: Mexico. Veracruz, Orizaba, Mueller2115 (holotype: LE; isotype: US-091892 fragm. ex LE!). Rhizomatous perennials, innovations extravaginal. Culms 120-180 cm tall, erect, scabrous; nodes 3-6. Leaf sheaths coriaceous, scabrous, brown, striate, older basal sheaths disintegrating into fibres; auricles absent; ligules 0.5-1 mm long, coriaceous, apex truncate; blades 35-70 x 0.4-1.2 cm, flat to involute at tip, green, abaxially scabrous, with prickles on adaxial and abaxial surfaces. Panicles 30-40 cm long, 25-40 cm wide, lax, pyramidal, branched; branches scabrous. Spikelets 12-15 mm long, lanceolate, florets 4 or 5; rachilla densely hairy; glumes 4.5-8 mm long, membranous, narrowly lanceolate, green, scabrous; lower glumes 4.5-6 mm long, 1-nerved; upper glumes 6-8 mm long, 3-nerved; lemmas 8-9 mm long, 5-nerved, membranous to coriaceous, lanceolate, green, scabrous and densely hairy, apex entire, awnless; callus 18 Festuca in South American Paramos sparsely hairy; paleas as long as adjacent lemma, membranous, papillose, keels scabrous; anthers about 2.9 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 4/5 as long as the grain. Leaf blade anatomy.•Cross sections with many vascular bundles; adaxial ribs small; sclerenchyma under both abaxial and adaxial epidermis, discontinuous, extending to the vascular bundles forming girders; adaxial epidermis with bulliform cells and without hairs. Observations.•Festuca amplissima is the only species represented in Festuca sect. Ruprechtia. It differs from other species of F. subg. Montanae by lacking cataphylls and by having glabrous ovaries. Distribution and habitat.•This species was confirmed in Mexico, Guatemala, Costa Rica, and Panama. Davidse et al. (1994) also mentioned this species from Venezuela. In Central America and Mexico this species is known from forests (Pinus spp. and Quercus spp.) and forest clearings. Additional specimens examined. COSTA RICA. Cartago: Canton de Turrialba, Parque Nacional Chirripo, Cuenca del Matina, estacion Crestones, sendero a Ventisqueros, 09°29'52"N, 83°29'20"W, 3400-3500 m, 11 May 1996, Gambo et al. 819 (MO). San Jose: Trail to the Valle de los Leones, lower part of Valle de los conejos, along the upper river Talari paramo formation with Chusquea sp. and areas of short burned forest, 83°31'W, 09°27'N, 3250-3450 m, 21-23 Aug 1971, Burger & Gomez 8246 (NY); Canton de Perez Zeledon, Parque Nacional Chirripo, cuenca Terraba - Sierpe Chirripo, base Crestones paramo, 8°27'25"N, 83°30'38"W, 3460 m, 12 Jul 1996, Alfaro 1027 (MO); Along the trail from Canaan to Chirripo via Los Angeles above N the Rio Talari, elfin forest of dead oak trees, 09°30N, 83°31 'W, 3200-3400 m, 19-22 Jan 1970, Burger etal. 7428 (F). GUATEMALA. Chimaltenango: Volcan de Agua, 10000 ft, 22 Jul 1937, JRJ 895 (F); Volcan de Acatenango, above Las Calderas, 2700-2900 m, 1 Mar 1939, Standley 61885 (F); above Las Calderas, 2700-2900 m, 1 Mar 1939, Standley 61899 (F). Sacatepequez: Volcan de Agua, above Santa Maria de Jesus, 2250-3000 m, 2 Nov 1939, Standley 65227 (F). MEXICO. Distrito Federal: Ajusco, Rancho Vieja Cuilotepec Tlalpan, 2730 m, 19 Sep 1985, E. Manriqu et al. 1147 (MO); Valle de Mexico, 2500 m, 1 Jan 1952, Matuda 25909 (MO). Chiapas: Mun. San Cristobal las Casas, at Piedracitas, 7 km E of San Cristobal along road to Zontehuitz, slopes with Quercus and Pinus, 7800 ft, 20 Jul 1965, D. Breedlove 11151 (F); Mun. Tenejapa, Paraje Matsab, 9000 ft, 5 Dec 1966, Alush Shilom Ton 957 (F). Guerrero: Mun. Heliodoro Castillo, ladera E del Cerro Teotepec, 68 km de Filo de Caballo por terraceria a El Paraiso, Bosque de Abies, Quercus, Pinus, Alnus y Clethra, 2730m, 23 Nov 1991, Gonzdlez-Ledesma etal. 484 (F); Mun. de Chilpancingo, Parque Omiltemi a 35 km W of Chilpancingo bosque mesofilo, 2450 m, 21 Nov 1991, Gonzdlez-Ledesma etal. 478 (MO). Jalisco: 14-18 km SW of Tequila on Volcan de Tequila Quercus, Arbutus forest, 2400-2600 m, 11 Jul 1974, D. Breedlove 39265 (MO). Michoacan: Mountain near Patzcuaro, 8500 ft, 23 Nov 1891, C.G. Pringle 3945 (NY); Mt. Tancitaro, rocky ledge, 10300 ft, 25 Jul 1941, Leavenworth etal. 1213 (F, MO,NY). Oaxaca: Distr. De Ixtlan, Mun. De Ixtlan, Rancho Teja, 0.25 km N of the upper casita, 2250 m, 4 Aug 1981, Martin 630 (US). Puebla: Mun. San Nicolas de los Ranchos 6 km al SE de Paso de Cortez, brecha a Xalitzintla, 20°18N, 98°44'W, 3400 m, 14 Sep 1988, Tenorio 15093 (US). Veracruz: Pico de Orizaba, 2500 m, Liebmann 6109, 6110 (C); Sep 1841, Liebmann 12904 (C). PANAMA. Chiriqui: Volcan Baru, in and near Potrero Muleto, abundant floor of crater 9400-10000 ft, 17 Nov 1978, B. Hammel 5619 (MO); Volcan Chiriqui, Boquete Distr., 10400 ft, 18 Jul 1938, Davidson 1043 (F, MO); W slope of Volcan Chiriqui, 11 km directly WNW of Boquete, ridge looking down on Potrero Muleto, elfin forest dominated by Ericaceae, 3000 m, 20 Nov 1975, G. Davidse &DArcy 10244 (NY). lb. Festuca amplissima subsp. sierrae Stancik, Preslia, Prague, 75: 341. 2003. (Fig. 1). TYPE: Colombia. Magdalena, Sierra Nevada de Santa Marta, Hoya del Rio Donachui, Cancurua, SE slopes, 2400-2650 m, 10 Oct 1959, J. Cuatrecasas & Castaneda 24736 (holotype: COL!; isotype: US!). This subspecies differs from typical form by having a larger, denser, and widely spreading panicles. Observations.•The outstanding character of larger panicles and the isolated geographical position in the northern Andes (Sierra Nevada de Santa Marta, Serranna de Perija), were the primary reasons for recognizing this subspecies. Distribution and habitat.•In Colombia and Venezuela Festuca amplissima subsp. sierrae Festuca in South American Paramos 19 100 x Figure 1. Festuca amplissima subsp. sierrae. A. Stylized growth form. B. Habit. C. Spikelet. D. Glumes. E. Lemma. F. Leaf blade cross-section. A-E, Cuatrecasas & Castaneda 24736 (COL). F, Barclay & Juajibioy 7033 (COL). occurs in matorral and shrub lands and the grass transitional zone of the paramo at an altitude of 2200-3400 m. Additional specimens examined. COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, alrededores de cabeceras de Rio Ancho, paramo de Macotama, 3490 m, 16 Feb 1959, Barclay & Juajibioy 7033 (COL, MO, US); Paramo de Macotama, NW side of valley, grassy slope, 3490 m, 13-18 Dec 1959, Barclay & Juajibioy 7017 (MO); Festuca in South American Paramos 20 Mun. Cienaga, Cabeceras del Rio Sevilla, 3500 m, Carbono 2474 (UTMC); transecto del Buritaca-filo La Cumbre, 3 020 m, 15 Aug 1977, Rangel et al. 945 (COL); about 30 mi inland from Dibulla, 3850 m, Jul 1923, Seifriz 489 (US). VENEZUELA. Zulia: Serrania de Perija, Serrania de los Motilones, Distr. Perija, headwaters of Rio Negro, 3000 m, 27 Jun 1974, Tillet & Honig 746-765 (MO). sheaths membranous, glabrous, striate, brown; auricles absent; ligules 0.5-0.7 mm long, membranous to coriaceous, apex truncate, ciliate; blades 15-25 cm long, 3-5 mm wide, flat, green, abaxially scabrous. Panicles 20-25 cm long, 2-5 cm wide, flexuous, pendant; branches scabrous. Spikelets 9-11 mm long, florets 3 or 4; rachilla ca. 1.2 mm long, pilose; glumes 2.5-6 mm long, narrowly lanceolate, coriaceous, purplish, pilose, apex acute; lower glumes 2.5-3.5 mm long, 1 -nerved; upper glumes 4.5-6 mm long, 3-nerved; lemmas 7.5-8.5 mm long, 5-nerved, lanceolate, membranous to coriaceous, margins short pilose, apex entire, awned or awnless, pilose, the awn 0-0.5 mm long; paleas as long as the lemma, pilose; lodicules ca. 0.8 mm long, oblong; anthers 2-2.5 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 4/5 of the total length. Leaf blade anatomy.•Cross-sections with numerous vascular bundles; adaxial ribs small, 2. Festuca coromotensis B. Briceno, Ernstia4 (3^t): 76. 1994. (Figs. 2,3,72E-F, 73A & B). TYPE: Venezuela. Merida, Dept. Libertador, Parque Nacional Sierra Nevada, Laguna La Coromoto, 3300 m, 3 Jul 1987, B. Briceno & G. Adamo 2003 (holotype: MERF!; isotype: MERC!). Loosely tufted, rhizomatous perennials with extravaginal innovations. Culms 100-120 cm tall, erect, scabrous; nodes 4 or 5 on lower 1/2. Leaf -80 -75 -70 $ f ' 10 in "X* ( <w PANAMA /r ^sC\j C* VENEZUELA COLOMBIA _^j. BRAZIL J^^^N- d • < *L f -80 ECUADOR A / ^w^" N 0 200 t 400 Kilometers ^ -75 Figure 2. Distribution of Festuca amplissima (•), F. coromotensis (A), and Fflacca (•). m f -70 ^ Festuca in South American Paramos 21 Figure 3. Festuca coromotensis. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 4173 (PRC). inconspicuous; sclerenchyma below abaxial and adaxial epidermis, discontinuous, forming girders; bulliform cells absent; abaxial epidermis with scattered prickles; adaxial epidermis with sparse hairs, the hairs ca. 0.1 mm long. Observations.•The habit of F. coromotensis is very similar to F. sodiroana. However, the spikelets of F sodiroana have shorter glumes (lower glumes 1.3-1.8 mm, upper glumes 2.2-3.5 mm), shorter lemmas (5-6.5 mm), and shorter anthers (0.8-1.2 mm). 22 Festuca in South American Paramos Distribution and habitat.•This species is endemic to Venezuela, known only from the Andean states of Merida and Tachira. Festuca coromotensis occurs in the Andean forest and matorral communities between 3000-3500 m. Additional specimens examined. VENEZUELA. Merida: Entre Los Arbolitos y Los Nevados, Paramo El Toro, 3100 m, 12 Oct 1988, B. Briceno et al. 2328 (Herbarium Briceno); Mun. Santo Domingo, Laguna Negra, patches of Polylepis, 08°47' 13.4"N, 70°48 '26.1 "W, 3500 m, 6 Nov 2000, D. Stancik 4173 (CAR, COL, PRC); Mun. Santo Domingo, Laguna Mucubaji, schrubby patches, 08°47'19.5"N, 70°48'36.1"W, 3470 m, 6 Nov 2000, D. Stancik 4174 (AAU, CAR, COL, PRC); Mun. Tabay, Laguna Coromoto, mountain forest, 08°35'41.7"N, 71°01 '26.9"W, 3100-3200 m, 7 Nov 2000, D. Stancik 4180 (AAU, CAR, COL, US, VEN, W); Mun. Tabay, Laguna Coromoto, mountain forest, 3000-3100 m, 7 Nov 2000, D. Stancik 4177B (PRC). Tachira: Paramo entre Enconejada para Pogonero, 3000 m, Sep 1955, Baldillo 3496 (VEN); Mun. La Grita, Par. La Negra, 8° 15' 11.2"N, 71°52'41.5"W, 3200 m, 10 Nov 2000, D. Stancik 4281 (AAU, CAR, COL, VEN); 8°15'26.4"N, 71°52'16.5"W, 3100 m, D. Stancik 4282 (CAR, COL, PRC). 3. Festuca elviae B. Briceno, Ernstia 4 (3^1): 77, f. 2-A. 1994. (Figs. 4,6,73C-F). TYPE: Venezuela. Merida, Libertador: Paramo La Culata, camino hacia Laguna Tapada, 3300 m, 19 Oct 1984, B. Briceno & G. Adamo 1114 (holotype: MERF!). Loosely tufted, rhizomatous, perennials with extravaginal innovations. Culms 40-80(-100) cm tall, erect, glabrous, nodes 2 or 3 in basal half. Leaf sheaths membranous, brown, glabrous; auricles absent; ligules 0.7-1.5 mm long, membranous, apex truncate, ciliate or dentate; blades 15-20 cm long, 1-6 mm wide, flat, sometimes involute, green, abaxially scabrous. Panicles 10-17 x 1-2 cm, contracted, erect; branches scabrous. Spikelets 7-9 mm long, florets 3-5; rachillas 1-1.4 mm long, glabrous; glumes 0.8-3 mm long, narrowly lanceolate, membranous, green, glabrous, apex acute; lower glumes 0.8-1.5 mm long, 1-nerved; upper glumes 2-3 mm long, 3-nerved; lemmas 4.5-5.5(-6) mm long, 5-nerved, lanceolate, membranaceous-coriaceous, green, apex glabrous or scabrous, entire, sometimes shortly awned, the awn 0.5-1 (-2) mm long; paleas almost as long as the lemma, apex scabrous; anthers 0.9-1.2 mm long; ovary apex glabrous. Caryopses oblong; hilum nearly the entire length. Leaf blade anatomy.•Cross sections with numerous vascular bundles and ribs on abaxial surface; sclerenchyma under both abaxial and adaxial epidermis discontinuous and extending to the vascular bundles; bulliform cells absent; abaxial epidermis with scattered prickles. Observations.•Spikelets with short glumes and short-awned lemmas suggest affinity with the Colombian and Ecuadorian species, F. andicola and F sodiroana. However, Festuca andicola and F sodiroana have slightly longer lower glumes (1.3-1.8 mm), nodding culms, and narrower panicles (0.4-1 cm). Distribution and habitat.•This species is endemic to Venezuela, known only from the state of Merida. It occurs in the humid or swampy margins of the Andean forest, along streams, in clearings, and in pastures, between 3000-3700 m. Additional specimens examined. VENEZUELA. Merida: Dept. Rangel, SE de la entrada a la Lag. Mucubaji, 3550 m, 28 Aug 1980, B. Briceno & Adamo 191, 196 (MERF, MO); 3550 m, 2 Oct 1996, B. Briceno et al. 3390 (Herbarium Breceno); Dept. Rangel, SE del Hotel Los Frailes, 3100 m, 22 May 1981, B. Briceno & Adamo 274 (MERF, MO); Dept. Rangel, camino hacia Laguna Las Canoas, 3100 m, 10 Jul 1981, B. Briceno & Adamo 306 (MERF); Paramo Los Granates, 3420 m, 30 Aug 1985, B. Briceno & Adamo 1320 (MERF); Paramo Misinta, 3100 m, 3 Dec 1986, B. Briceno 6 Adamo 1705 (MERF); Mun. Santo Domingo, Laguna Mucubaji, swamps around the lagoon, 08°47'33"N, 70°48'56"W, 3600 m, 6 Nov 2000, D. Stancik4171 (AAU, CAR, COL, PRC); Laguna Mucubaji, shrubby patches, 3470 m, 8°47'20"N, 70°48'36"W, D. Stancik 4175 (AAU, CAR, COL, PRC). Mun. Mucuchies, Paramo de Piedras Blancas, 8°48 '31"N, 70°55 '08"W, 3700 m, 4 Nov 2000, D. Stancik 4215 (AAU, CAR, COL, PRC); Mun. La Culata, Paramo La Culata, 8°45'N, 71°33'W, 3300 m, 12 Nov 2000, D. Stancik4250 (CAR, COL, PRC); 3100 m, D. Stancik4257 (CAR, COL, PRC); 8°45 'N, 71 °03' W, 3100 m, 12 Nov 2000, D. Stancik 4256 (CAR, COL, PRC, US); Dept. Libertador, NE con respecto al Valle de la Culata., Dept. Libertador, paramo La Culata, Laguna Tapada, 3300 m, 7 Jun 1984, B. Briceno & Adamo 948 (Herbarium Briceno), 28 Sep 1984, B. Briceno & Adamo 1086 (MERF); Pass on the Merida-Barinas Hwy, paramo Festuca in South American Paramos 23 2 cm 1 mm Figure 4. Festuca elviae. A. Habit. B. Inflorescence (detail). C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Briceno & Adamo 948 (PRC). 24 Festuca in South American Paramos above Laguna Grande et Universidad de los Andes, 3500m, 19Nov 1971, G. Davidseetal. 3223(MO); Valle de los Calderones y Paramo de los Pozones, 3400 m, 19 Oct 1995, B. Briceno et al. 3345 (herbarium Briceno); Laguna Coromoto, mountain forest, 8°35'41"N, 71°01'27"W, 3000-3100 m, 7 Nov 2000, D. Stancik 4178 (AAU, CAR, COL, PRC, US, W); Entre La Mucuy e Laguna Coromoto, 3000 m, 27 Mar 1994, B. Briceno et al. 2601 (herbarium Briceno). 4. Festuca flacca Hack, ex E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 69: 1543. 1984. (Figs. 2, 5, 74A-D). TYPE: Ecuador. Pichincha, in silv. super, pr. Tablahungi, 1887, Sodiro 36/11 (holotype: W!; isotypes: QPLS!, W!,US!). Festuca subulata var.fraseriana St.-Yves, Candollea 3:451.1928. TYPE: Amerique du Sud, Equateur, sin. loc, 1860, Fraser s. n. (holotype: G!). Loosely tufted perennials with extravaginal innovations. Culms 70-120(-150) cm tall, erect, glabrous; nodes 3-7 nodes in distal half. Leaf sheaths membranous, brown, striate, scabrous, occassionally hairy, margins free; auricles absent; ligules 3-3.5 mm long, membranous, apex acute; blades 25-30 x 0.3-1.1 cm, flat, green, abaxially scabrous. Panicles 20-35 x 5-15 cm, lax, fewflowered, linear-oblong to ovate; branches scabrous. Spikelets 7-8 mm long, florets 2-4 with a rudiment; rachilla pilose; glumes 1.2-3.5 mm long, green, glabrous, narrowly lanceolate, apex acuminate; lower glumes 1.2-2.4 mm long, 1-nerved; upper glumes 2.5-3.5 mm long, 3-nerved, sometimes scabrous on dorsally; lemmas 6.5-7 mm long, 5-nerved, membranous to coriaceous, lanceolate, green, occasionally purplish-green, papillose, apex scabrous, awned, the awn 9-12 mm long, scabrous, fine and straight; paleas almost as long as the lemma, glabrous and inconspicuously scabrous on margins and keels, apex hairy; lodicules lanceolate, acuminate; anthers 1.1-1.4 mm long; ovary apex sparsely hairy. Caryopses lanceolate; hilum 4/5 the length. Leaf blade anatomy.•Cross sections with numerous vascular bundles and without ribs above; sclerenchyma under both abaxial and adaxial epidermis, discontinuous, extending to the vascular bundles forming girders; bulliform cells present; abaxial epidermis (or both sides) with scattered prickles. Observations.•Alexeev (1984) incorrectly cited the page of the original publication of Sodiro (1889), where this taxon is mentioned for the first time then later described by Alexeev as F flacca. Festuca flacca is morphologically similar to other long-awned species that have short glumes, such as: F ulochaeta from Brazil, Colombia, and Venezuela; F. cochabambana from Bolivia; and, F tovariensis from Bolivia and Peru. However, Festuca ulochaeta has flexuous lemma awns, F tovariensis has truncate ligules, and F cochabambana has pubescent leaf sheaths. Distribution and habitat.•Festuca flacca is endemic to Ecuador, known from the central and northern Andean departments (Carchi, Cotopaxi, Imbabura and Pichincha). It occurs in mountain Andean forests between 2900-3500 m. Additional specimens examined. ECUADOR. Carchi: Hacienda San Rafael, faldas occidentales de la Cordillera Oriental, 2900-3000 m, AcostaSolis 21026 (US); wooded hills about 5 mi S of Tulcan, 2500 m, A.S. Hitchcock 21042 (US). Cotopaxi: Parque Nacional Cotopaxi, pine plantation, 00°37'S, 78°27'W, 3400 m, S. Laegaard 54540A (AAU, QCA), S. Laegaard 54540B (AAU, QCA). Cotopaxi, 3550 m, E. Asplund 6359 (NY, S, US). Imbabura: Lake Cuicocha - Islote Chica, 3150 m, E. Asplund 7156 (NY, S, US); Laguna Mojanda, 10 km SSW of Otavalo, 00°10'N, 78°18'W, 2900-3150 m, Sparre 13562 (S). Pichincha: Road Pifo-Papallacta, km 15, Polylepis incana forest, 00°16'S, 78°17'W, 3530 m, 16 Apr 1992, S. Laegaard 102316 (AAU, QCA, QCNE); Pifo, 3000 m, Mille s.n. (QPLS); Volcan Pasachoa, above house of Fundacion Natura, 00°27'S, 78°30'W, 2900-3300 m, 16 Sep 1985, S. Laegaard 55260 (AAU, QCA, QCNE); 3200 m, 27 Apr 1985, S. Laegaard 54164 (AAU, QCA, QCNE); 19 Sep 1985, S. Laegaard 55281 (AAU, QCA, QCNE); Mt. Pichincha - St. Gertriadis, Sodiro s.n. (QPLS); Mt. Pichincha Tablahuras, Sodiro s.n. (QPLS); Mt. Pichincha, Sodiro s.n. (US); Slope of Pichincha above Lloa, 3200 m, E. Asplund 7535 (AAU); E. Asplund 7559 (K, P, S,US); 2900-3100 m, Acosta-Solis 20054 (US); Concepcion, near Hda. Monjas, 10200 ft, Bell 17 (S); Mun. Amaguana, Volcan Pasochoa, bosque andino, 3400 m, 14 Sep 2000, D. Stancik 3668 (PRC, QCA, US); Mun. Otavalo, road from Laguna Mojanda to Cochasqui, shrubby margin of the road, 00°04'55.1"N, 78°17'49.8"W, 3450 m, 19 Oct 2000, D. Stancik 4104 (PRC, QCA). Festuca in South American Paramos 25 2 cm Figure 5. Festuca flacca. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 3668 (PRC). Festuca in South American Paramos 26 -80 -75 -65 -70 i, •• -60 0 m. PANAMA 0 ^JOOA^HCIL 10 4>W L? ^^•^ *^^& 1 #^ \ 10 VENEZUELA -^ • ] • ^-rl • r % COLOMBIA { L> \ W BRAZIL f J# ECUADOR .T "1/ -80 / 1 0 -75 N 200 -70 400 Kilometers -65 li -60 Figure 6. Distribution of Festuca sodiroana (•), F. elviae (A), and F. guaramacalana (•). 5. Festuca guaramacalana Stancik, Novon 14(3): 343. 2004. (Figs. 6, 7, 74E-F, 75A & B). TYPE: Venezuela. Trujillo, Mun. Bocono, Parque Nacional Guaramacal, 09°14'15"N, 70° 11' 14"W, Andean mountain forest, margin of the brook with Neurolepis sp., Cortaderia sp., 2880 m, 29 Nov 2000, D. Stancik 4286 (holotype: PRC!; isotypes: CAR!, COL!). Rhizomatous perennial, forming small tussocks with extravaginal innovations. Culms 100• 130 cm tall, erect, glabrous; nodes 3 or 4 nodes in distal half. Leaf sheaths membranous-coriaceous, purplish-brown, striate, fibrous at base, margins free; auricles absent; ligules 1-1.5 mm long, membranous, sometimes coriaceous, apex acute; blades 30-40 x 0.6-0.7 cm, conduplicate to involute, green, sometimes olive-green, abaxially glabrous. Panicles 15-20 x 2-5 cm, flexuous, pendant, elon- gate, branched; branches finely scabrous. Spikelets 13-15 mm long, narrowly lanceolate, florets 5-7; rachilla pilose; glumes 3.7-7 mm long, narrowly lanceolate, purplish, membranous to coriaceous, sparsely scabrous, apex acute; lower glumes 3.74.7 mm long, 1-nerved; upper glumes 5-7 mm long, 3-nerved; lemmas 9.5-10 mm long, membranous to coriaceous, lanceolate, 5-nerved, purplish, papillose, apex entire, mucronate or shortly awned, the awn 0.5-1 mm long; callus sparsely pilose; paleas finely pilose, almost as long as the lemma; lodicules ovate, two-dentate; anthers 2.5-2.8 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 4/5 as long as the grain. Leaf blade anatomy.•Cross-sections with about 11 vascular bundles and 5 ribs above; sclerenchyma under both abaxial and adaxial epidermis discontinuous and extending to some vascular bundles forming girders; bulliform cells absent; Festuca in South American Paramos 27 Figure 7. Festuca guaramacalana. A. Growing form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 4286 (PRC). 28 Festuca in South American Paramos adaxial epidermis with scattered microhairs, ca. 0.09 mm long. Observations.•Festuca guaramacalana belongs to Festuca sect. Subulatae and among the South American members of this section, F. guaramacalana has the largest spikelets and longest glumes (Stancik & Peterson 2002). The lemmatal awns of F. guaramacalana are short (0.5-1 mm) and this is in contrast with most other members of sect. Subulatae, although F. sodiroana typically has acute lemmas. Morphologically, F. coromotensis appears to be most similar to F guaramacalana but the former has shorter ligules (0.5-0.7 mm long), ligules with truncate apices, shorter spikelets (9-11 mm long) with 3 to 4 florets, and shorter lemmas (7.5-8.5 mm long). Distribution and habitat.•Festuca guaramacalana is endemic to Venezuela and is known only from Parque Nacional Guaramacal where it occurs near small brooks with Neurolepis sp. and Cortaderia sp. between 2600-2880 m. This species is a narrow endemic and is vulnerable to extinction due to loss of habitat. Additional specimens examined. VENEZUELA. Trujillo: Mun. Bocono. Parque Nacional Guaramacal. Paramo El Pumar. 2870 m, 2004, M. Ramirez & N. Cuello 3523, 3524, 3528 (PORT). 6. Festuca sodiroana Hack, ex E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 69: 1545. 1984. (Figs. 6, 8, 75C-F). TYPE: Ecuador. Pichincha, silv. super, et pasq., 1884, Sodiro 36/6 (holotype: W!; isotypes: MO-923640!, PRC ex QPLS!, QPLS!, US!). Rhizomatous perennials with extravaginal innovations. Culms (40-)50-120(-150) cm tall, erect, glabrous; nodes 2 or 3(-^t) in distal half. Leaf sheaths membranous, brown, striate, pilose at base, margins free; auricles absent; ligules 0.2-1.1 mm long membranous to coriaceous, apex truncate, ciliate; blades 15-25 x 0.4-0.7 cm, flat, green, abaxially glabrous. Panicles 15-20 x 1-2 cm, lax, contracted and something nutant, branched; branches scabrous. Spikelets 7.5-9(-10) mm long, florets 4 or 5(-6); rachilla pilose; glumes 1.3-2.9(-3.5) mm long, lanceolate, green, membranous to coriaceous, glabrous, apex acute; lower glumes 1.3-1.8 mm long, 1-nerved; upper glumes 2.2-2.9(-3.5) mm long, 3-nerved; lemmas 5-6(-6.5) mm long, 5-nerved, membranous to coriaceous, lanceolate, green, awnless, apex acute, entire, muticous, scabrous; paleas almost as long as the lemma, glabrous or inconspicuously scabrous, apex hairy; lodicules lanceolate, acuminate; anthers 0.8-1.2 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 4/5 of total length. Leaf blade anatomy.•Cross-sections with numerous vascular bundles, without ribs above; sclerenchyma under both abaxial and adaxial epidermis, discontinuous, extending to the vascular bundles; bulliform cells present; abaxial epidermis with scattered prickles, adaxial epidermis without hairs or with scattered hairs. Observations.•Specimens that belong to this species were originally determined by Hackel and Sodiro as either F. sodiroana or F pichinchae, but these names were not validly published. Alexeev recognized that these specimens belonged to the same species and validated the name F sodiroana. Festuca sodiroana differs from others in sect. Subulatae by having muticous lemmas. Distribution and habitat.•Known from southern Ecuador to northern Colombia where it is found in all three Cordilleras. It occurs in forest clearings, margins of brooks in Andean mountain forests, roads banks, and trails at an altitude of 2600-3800 m. Festuca sodiroana is known to occur in vegetation communities: Neurolepido aristatae-Oreopanacion nitidii (Cleef et al. 1983) and Chusquea scandentis-Weinmannion rollottii (Cleef etal. 1983). Additional specimens examined. COLOMBIA. Antioquia: Mun. Urrao, Paramo Frontino, Llano Grande, 3520 m, 10 Sep 1986, Rolddn etal. 360 (HUA); Campanas, La Laguna, 3500-3800 m, 3 Mar 1989, MacDougal et al. 4504 (HUA, MO). Cauca: Macizo Colombiano, Paramo de Las Papas, entre Boqueron y La Hoya, 2910 m, 11 Sep 1958, Idrobo et al. 3026 (COL); Parque Nacional Purace, termales de San Juan, 3100-3300 m, 6 Apr 1985, Wood 4792 (COL, K); Paramo de Moras, between Mosoco y Pitayo, 3000-3500 m, Feb 1906, Pittier 1511 (US). Mt. Purace, 3100-3300 m, 16 Jun 1922, Killip 6709 (US). Cundinamarca: Paramo de Tablazo, 3200 m, 8 Apr 1984, Wood 4347 (COL, FMB, K). Nariflo: El Encano, road from Pasto to Virgen, vereda Caltapamba, 3100 m, 23 Mar 1999, D. Stancik 2992 (COL, PRC, PSO, US); D. Stancik 2991 (COL, PRC, PSO); El Encano, road to village Colon, km 4, 2650 m, 13 Mar 1999, D. Stancik 2866 (COL, PRC, PSO); km 6, 2900 m, 13 Mar 1999, D. Stancik 2858 (COL, PRC, PSO); Mun. Festuca in South American Paramos 29 100 x Figure 8. Festuca sodiroana. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 2992 (PRC). 30 Festuca in South American Paramos Cumbal, vereda Las Huertas, 3600 m, 9 Mar 1999, D. Stancik 2754 (COL, PRC, PSO); Cumbal, lakeshore, 4000 m, 24 Mar 1941, Sneider 430 (NY); Mun. Pasto, paramo Puerto Frio, between villages Las Almas and Alisales, 2900 m,14 Mar 1999, D. Stancik 2870 (COL, PRC, PSO); Mun. Tuquerres, Volcan Azufral, road from vereda San Roque Alto, 3500 m, 9 Mar 1999, D. Stancik 2772 (COL, PRC, PSO); 2650m, 9Mar 1999,/). Stancik2776(COL, PRC, PSO); Vereda El Carmelo and La Florida, Volcan Dona Juana, 2900 m, 18 Mar 1999, D. Stancik 2902 (COL, PRC, PSO); Paramo de Bordoncillo, Mun. Santiago, vereda San Antonio de Bellavista, 1°11'N, 77°06'W, 3200-3400 m, 18 Mar 1964, Alberto et al. 144 (COL); Mun. Guachucal, paramo de Infernillo, 3200 m, 5 Mar 1999, D. Stancik 2632 (COL, PRC). Norte de Santander: between Pamplona and Mutiscua, 2700 m, 1 Apr 1984, Wood 4326 (COL, FMB, K). Putumayo: Paramo de Tabano, 15 May 1935, Archer 3402 (US). Santander: Valle California, 2800 m, 14 Sep 1985, Wood5066 (FMB, K). Tolima: Mun. Ibague, Nevado del Tolima, 04°37.4'N, 75°19.8'W, 3400 m, 8 Jun 2000, D. Stancik 3595 (COL, PRC, PSO); 3400 m, 16 Dec 1984, Wood4648 (COL, FMB, K); W slope of Paramo Rico, 3200 m, 15-19 Jan 1927, Killip & Smith 17861 (NY), sin. loc: Cerro Nevada, Bos. La Pena, 2900 m, Lindig 1116 (K, P); Lindig 1117 (P); Mutis 5544 (US). ECUADOR. Azuay: Road Cuenca - Saraguro, km 6, S of Cumbe, 03°04'S, 79WW, 3150 m, S. Laegaard 105132 (AAU); Parque Nacional Cajas-Laguna Llaviuco, 3150 m, 22 Apr 1990, P.M. Peterson 8872, C.R. Annable & M.E. Poston (K, MO, QCA, QCNE, US); Ganadel, 3250 m, E. Asplund 17846 (S); between Cuenca and Hiugra, 2700-3000 m, A.S. Hitchcock 21682 (US); road Sigsig-Guadaquiza, 02°09'S, 78°43 'W, 3300 m, 29 May 1992, S. Laegaard et al. 103032 (AAU, QCA, QCNE); Mun. Cuenca, Parque Nacional Cajas, Laguna Llaviucu, 02°50'30"S, 79°08 '45"W, 3100-3150 m, 3 Aug 1999, Police 1 (PRC); Police 3 (PRC); Police 4 (PRC). Bolivar: Road Guaranda-Pueblo Viejo, km 21.6, 01°35'S, 79°09'W, 2500 m, 6 Mar 1988, S. Laegaard & S.A. Renvoize 70580 (AAU); km 20.1, 01°35'S, 79°05'W, 2700 m, S. Laegaard & S.A. Renvoize 70577 (AAU, K, QCA, QCNE). Caflar: Road Canar-Biblian, Fuganillas, 02°36'S, 78°54'W, S. Laegaard 52754 (AAU, QCA, QCNE); Tipococha-Hacienda Shical, 3000 m, Acosta-Solis 16963 (US); Acosta-Solis 16974 (US). Carchi: El Angel, Hacienda la Esperanza, 00°39N, 77°54'W, 3300 m, S. Laegaard 53118 (AAU, QCA); El Voladero, 00°38'N, 67°53'W, 3400-3800 m, Davalos 22 (US); wooded hills about 5 mi S of Tulcan, 2500 m, A.S. Hitchcock21094 (US); Road Las Juntas-El Angel, 00°46N, 77°46'W, 3180 m, 11 Mar 1992, S. Laegaard 101711 (AAU, QCA, QCNE); La Rinconada, 3200 m, E. Asplund 7195 (F, S); 3000 m, A.S. Hitchcock 20798 (NY, US). Cotopaxi: Road Pilalo-Zumbagua, 15 km above Pilalo, 00°59'S, 78°58'W, 3350 m, Holm-Nielsen 24604 (AAU); km 9, 0°59'S, 78°57'W, 3150 m, 7 Apr 1992, S. Laegaard 102227 (AAU, QCA, QCNE); RoadPilalo-Latacunga, 00°57'S, 78°58'W, 3400 m, Holm-Nielsen 1482 (AAU, MO, S); Road Quevedo-Latacunga, 3600 m, Hading et al. 8904 (GB); Rio Chalupas, 00°50'S, 78°21'W, 3700 m, S. Laegaard 101781 (AAU, QCA, QCNE); Mun. Lasso, Volcan Cotopaxi, margin of the forest at the road to National Park entrance, 00°39'6"S, 78°30'55"W, 3530 m, 28 Sep 2000, D. Stancik 3885, 3886 (PRC, QCA). Chimborazo: Road Chunchi-Zhub, km 22,02°22'S, 78°57'W, 2780 m, 27 May 1992, S Laegaard et al. 103010 (AAU, QCA); Urbina towards Mt. Chimborazo, 3750 m, E. Asplund 7878 (S); Pallatanga-comunidad Jesus del Gran Poder, 01°58'S, 78°56'W, 2800-3200 m, Clark et al. 1372 (QCNE). Imbabura: Road Otavalo-Selva Alegre, km 25.2,00°16'S, 78°24'W, 3300 m, S. Laegaard et al. 70813 (AAU, K, QCA); Paramo de Angochagua, 2900-3600 m, AcostaSolis 18839 (US); Cayambe-Laguna San Marcos, 11200 ft, Cazalet & Pennington 5417 (K, NY, US); Mun. Urcuqui, road to Cerro Yanaurcu, 00°26'28"N, 78° 15 '24"W, 4100 m, 15 Oct 2000, D. Stancik 4095 (PRC, QCA); Mun. Otavalo, road from Laguna Mojanda to Cochasqui, 00°04'55"N, 78°17'50"W, 3450 m, 19 Oct 2000, D. Stancik 4105 (PRC, QCA); D. Stancik 4113 (PRC, QCA). Loja: Road to Fierra Urcu, 03°33 'S, 79' 15'W, 3100 m, S. Laegaard et al. 18855 (AAU, LOJA, QCA, QCNE); Road Saraguro-Yacuambi, 03°34'S, 79°06'W, 3050 m, S. Laegaard 20688 (AAU, LOJA); between San Lucas and Ona, 2200-3100 m, A.S. Hitchcock 21512 (US); Udo de Sabanilla, 04°27'S, 79°09'W, 2700 m, 1 Sep 1998, S Laegaard et al. 19095 (LOJA); Mun. Saraguro, road to Fierra Urcu, 03°42'40"S, 79°18'12"W, 3400-3450 m, 24 Aug 2000, D. Stancik 3771, 3781 (PRC, QCA); 30003100 m, D. Stancik 3792 (PRC, QCA). MoronaSantiago: road Gualaceo-Limon, km 12 of pass, 03°01'S, 78°37'W, 2590 m, S. Laegaard et al. 103079 (AAU); Mun. Alao, way from Alao to Festuca in South American Paramos Parque Nacional Sangay, 3500-3700 m, 23 Jul 1999, D. Stancik 3292 (PRC, QCA, W); Parque Nacional Sangay, patches of the forest above the Quebrada Tablamitza, 3600 m, 22 Jul 1999, D. Stancik 3352 (PRC, QCA). Napo: km 45 on road Salcedo-Napo, 6 km NE, 00°56'S, 78°23'W, 3600 m, S. Laegaard 53372 (AAU, QCA, QCNE). Pichincha: Near Quito, Jun 1922, Harteman 39 (US); Paramo Guamani, 00°15'S, 78°12'W, 3700 m, S. Laegaard 105069 (AAU); 3530 m, S. Laegaard 102318 (AAU, QCA, QCNE); Volcan Pichincha, 3250 m, Asplund 6155 (AAU, MO); above Lloa, 3200 m, E. Asplund 7537 (F, S); 3250 m, E. Asplund 6155 (F, MO, S, US); Mt. Pichincha, Sodiro s.n. (NY, MO); 3400 m, Sodiro s.n. (US); Milles et al. 281 (US); Sodiro s.n. (US); 3600 m, Sodiro 282 (US); 3300 m, E. Asplund 16163 (S); At base of Mt. Pichincha-La Carolina, Sodiro s.n. (S); road Chillogallo-Chiriboga, km 3 E of San Juan, 3100 m, Sparre 16950 (S); Sparre 16920 (S); San Juan towards Quito, 3400 m, E. Asplund 16094 (S); Tambillo, 2700 m, E. Asplund 6215 (AAU, F, MO, S, US); road Pifo-Pintag, 00°19'S, 78°17'W, 3100 m, S. Laegaard 102293 (AAU, QCA, QCNE); road Santo Domingo-Quito, between Saloya and Chiriboga, 2800 m, Hurling et al. 10415 (GB); Volcan Corazon, 11000 ft, Prescott 913 (NY); road km 13.5 ESE of Machachi, and 10 kmNE towards Sangolqui, 3350 m, 25 May 1990, P.M. Peterson 9313 & E.J. Judziewicz (K, MO, QCA, QCNE, US); road Quito-Nono, 00°06'S, 78°31 'W, 3300 m, 17 Jun 1984, S. Laegaard 52285 (AAU, QCA, QCNE); Nono, 2600 m, E. Asplund 7462 (S); Volcan Pasochoa, 00°21 'S, 78°29'W, 3100-3400 m, 23 Feb 1992, S. Laegaard 101406 (AAU, QCA); 00°27'S, 78°30'W, 3200 m, S. Laegaard 55282 (AAU, QCA, QCNE); Sodiro s.n. (QPLS); Highway Aloag-St. Domingo, 2900 m, Sparre 15076 (S); Mun. Pifo, Paramo Guamani, Polylepis forest, 00°19'S, 78°15'W, 3700m, 19 Jun 1999,D. Stancik 3010 (PRC, QCA); Mun. Amaguana, volcan Pasachoa, 3400 m, 14 Sep 2000, D. Stancik 3667 (PRC, QCA); Mun. San Antonio, Res. Geobotanica Pululahua, 00°02'20"N, 78°29'33"W, bottom of the crater, 2700 m, 13 Sep 2000, D. Stancik 3656 (PRC, QCA). Tungurahua: sin loc, Spruce 5938 (K, W). Mun. Banos, Volcan Tungurahua, on way to refugium, E side of the volcan, 3300 m, 29 Jul 1999, Police 9 (PRC); Mun. Pillaro, Las Llanganatis, 01° 09'43"S, 78°15'9"W, 3600 m, 28 Sep 2000, D. Stancik 3985 (PRC, QCA). Zamora-Chinchipe: road Loja-Zamora, ca. 2-6 km E of pass, 03°59'S, 31 79°09'W, 2750 m, S. Laegaard 18748 (AAU, LOJA); 2600 m, S Laegaard 18727 (AAU, LOJA); Road Loja-Zamora, 1 km E of pass, 04°00'S, 79°09'W, 2700 m, S. Laegaard 18483 (AAU, LOJA, QCA, QCNE). 7. Festuca tovariensis Stancik & P.M. Peterson, Sida 20(1): 24.2002. (Figs. 9,11). TYPE: Peru. Huancavelica, Pro v. Tayacaja, Chuspi-Hda. Tocas, entre Colchabamba y Paucarbamba, monte bajo, 2800 m, 22 Apr 1954, O. Tovar Serpa 2057 (holotype: US-2181286!; isotype: USM!). Loosely tufted perennials with extravaginal innovations. Culms 70-90 cm tall, erect, glabrous; nodes 2•4 nodes in basal half. Leaf sheaths membranous, brown, margins free; auricles absent; ligules 1-2 mm long, membranous, margins ciliate, apex truncate; blades 8-15 cm long, 1.5•4.5 mm wide, flat, green, abaxially scabrous with ribs on abaxial surface. Panicles 15-20 x 7-10 cm, open, pendant; branches scabrous. Spikelets 7.5-9.5 mm long, florets 3; rachillas 1.1-1.4 mm long, densely pilose; glumes 1.5•4.5(•5) mm long, narrowly lanceolate, coriaceous, purplish, glabrous, apex acute sometimes scabrous; lower glumes 1.5-1.8 mm long, 1-nerved; upper glumes 3.5-4.5(-5) mm long, 1-3-nerved; lemmas 5.5-6.5 mm long, lanceolate, coriaceous to membranous, inconspicuously 5-nerved, purplish-green, scabrous, apex entire, awned, the awn 5-7 mm long, terminal, scabrous, straight; paleas as long as the lemma, keels scabrous, apex hairy; lodicules ca. 0.8 mm long, lanceolate, acuminate; anthers 1.5-1.6 mm long; ovary apex sparsely hairy. Caryopses lanceolate; hilum 2/5-1/2 of total length. Observations.• Festuca tovariensis is morphologically similar to other long-awned species that have short glumes, such as: F ulochaeta from Brazil, Colombia, and Venezuela; F cochabambana from Bolivia; and Fflacca from Ecuador. Distribution and habitat.•This species is known only from the Andean forest zone of northern and central Peru between 2500-3850 m. Additional specimens examined. PERU. Ancash: Pro v. Yungay, Parque Nacional Huascaran, Llanganuco sector, Maria Josefa trail between Chinancocha and Pucayacu, 09°05'S, 77°39'W, 3700-3850 m, 5 Jul 1985, Smith 10561 (MO). Ayacucho: Pro v. Huanta/La Mar, Tambo, WolkenNebel-Buschwald, 37 km to Ayna, 3250 m, 23 Mar 1977, Ellenberg 7024 (USM). Cajamarca. Prov. 32 Festuca in South American Paramos 2 mm Figure 9. Festuca tovariensis. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Caryopsis. A-H, Vega et al. 1544 (F). Festuca in South American Paramos Cajamarca, Yumagual, entre San Juan y El GavilanGavilan, ladera con arbustos, 2500 m, 6 Oct 1975, Vega etal. 1544 (F); Prov. Contumaza, alrededores de Guzmango, ladera, 2600 m, 25 Jul 1992, Sagdsteguii 14785 (F); Carretera a Yumagual, 2500 m, 26 Jun 1966, Vega 249 (USM). 8. Festuca ulochaeta Nees ex Steud., Syn. PI. Glumac. 1:305.1854. (Figs. 10,11,76A-D). TYPE Brazil. Sellow s.n. (isotypes: B!, K!) Festuca leptothrix Trin. ex Doll, Fl. Bras. 2(3) 115. 1878. TYPE: Brazil. Sao Paulo, G.H. von Langsdorffs.n. (holotype: LE-TRIN-2818.01! isotypes: K!, US-91399 fragm!). Vulpia ulochaeta Nees ex Doll, Fl. Bras. 2(3): 115 1878, nom. inval. TYPE: Brazil. Sellow s.n (holotype: B!, BAA-3489 fragm ex B). Loosely tufted perennials with extravaginal innovations. Culms 60-120 cm tall, erect, glabrous; nodes 2-4 nodes. Leaf sheaths membranous, brown, striate, glabrous; auricles absent; ligules 0.5-1 (-2) mm long, membranous, apex truncate; blades 15-30 x 0.5-1.1 cm, flats, green, abaxially scabrous. Panicles 15•25(•35) x 15-25 cm, flexuous, pendant, ovate; branches scabrous. Spikelets 9-12 mm long, florets 3-5; rachilla shortly hairy; glumes 2.5-4.5(-6) mm long, membranous to coriaceous, narrowly lanceolate, green, apex acute, upper third scabrous; lower glumes 2.5-3.5(-4) mm long, 1 -nerved; upper glumes 3.5-4.5(-6) mm long, 3-nerved; lemmas 6-8(-9) mm long, 5-nerved, membranous to coriaceous, lanceolate, green, papillose, apex scabrous, awned, the awn 7-15 mm long, scabrous, flexuous; paleas almost as long as the lemma, glabrous, margins and upper third scabrous; lodicules lanceolate; anthers 1.1-1.5(-2) mm long, ovary apex sparsely hairy. Caryopses lanceolate; hilum 3/4 of total length. Leaf blade anatomy.•Cross-sections with numerous vascular bundles, without ribs above; sclerenchyma under both abaxial and adaxial epidermis, discontinuous and extending to thevascular bundles; bulliform cells absent; abaxial epidermis with scattered prickles. Observations.•Festuca ulochaeta has spikelets with short glumes and long-awned lemmas, characteristics shared with Fflacca, F cuzcoensis, and F cochabambana. Festuca ulochaeta differs from these other species by having lemmas with extremely long (7-15 mm) and distinctly flexuous (versus straight) awns. 33 Distribution and habitat.•Festuca ulochaeta is known from the humid forests of SE Brazil and NE Argentina between 700-2200 m. In Colombia and Venezuela this species occurs in clearings, margins of the streams, and roadsides in Andean forests (Cordillera Oriental and Aragua, Merida) between 2600-3100 m. Festuca ulochaeta is reported from Costa Rica for the first time here. Additional specimens examined. ARGENTINA. Misiones: Dpto. Gral. Manuel Belgrano, Santo Andresito,26°12'S,53°40'W, 720 m, 15 Feb 1996, O. Morrone & A.M. Cialdella 854 (CTES, MO). Salta: Dpto. Santa Victoria, camino de Toldos a Lipeo, a 15 km de Toldos, 1650 m, 11 Nov 1974, A.M. Turpe 2932 (W); Arroyo Latas, 20 Feb 1924, L. Parodi 5673 (US). BRAZIL. Minas Gerais: Xanxere, Pinheiral, 9 km E of Xanxere, 600-800 m, 26 Feb 1957, Smith & Klein 11836 (B, R, US); Serra de Caparao, 2100-2220 m, 4 May 1925, A. Chase 9659 (W); 2100 m, 4 May 1925, A. Chase 9672 (US); 2100-2200 m, A. Chase 9673 (US); Monte Verde, Sep 1997, Wagner 5010 (ICN); Vila Monteverde, Pico da Pedra Salada, Burman 880 (SP). Parana: ca. 85 km of Guarapuava, 800-1050 m, 6 Mar 1967, Linderman et al. 4658 (K, W, US); Ypiranga, 9 Feb 1904, Dusen 3624 (R); Dusen 3515 (R); Piraquara-Estrada a Monte Algre, Apr 1950, G. Hatschbach 1912 (BAA); Tres Barras, 27 Jan 1916, Dusen 17561 (BAA); Iraty, in silvula subuliginosa, 26 Feb 1909, Dusen 7808 (US); Curitiba, woods along stream, Estacao Experimental, 13 Feb 1946, J.R. Swollen 8540 (US); Curitiba, colonia Muricy-S. Jose dos Pinhais, na beira da Estrada, 1 Mar 1965, Scito & Kuniyoshi 1315 (K); Curitiba, Orladobrejo, 15 Jan 1975, Maguire 3487 (K);Orla Guarapuava, fazenda Capao Redondo, 20-23 Mar 1946, J.R. Swollen 8859 (US); Mun. Palmas, 1100 m, 16 Feb 1958, G. Hatschbach 4715 (US); Curitiba, Parque Barigui, area degradada, proxima a orla da foresta con Araucaria, 25°22'S, 49°13'W, 2 Apr 1997, Kozera & Izernhagen 476 (NY); Piraguara, ad marginam, 1 Jul 1909, Dusen 7783 (K, NY); Mun. Rio Branco do Sul, Dombrowski 2432 (ICN, K); Sao Mateus do Sul, Rio Potinga, 760 m, do interior da mata, 9 Feb 1966, G. Hatschbach 13820 (K). Rio de Janeiro: Serra de Itatiaia, 1960 m, 16 Jan 1925, A. Chase 8276 (RB, US); Parque Nacional de Itatiaia, entre Agulhas Negras e Abrigo Massena, campo alto 11-12 Feb 1985, Burman 905 (SP). Rio Grande do Sul: Highway BR-116 to Lajes, 1000 m, 10 Mar 1976, G. Davidse et al. 11074 (K, SP, VEN); Entre J. Kroeff et Roncinka, 18 Mar 1964, 34 1 mm Festuca in South American Paramos 1 mm Figure 10. Festuca ulochaeta. A. Stylized growth form. B. Habit. C. Inflorescence. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 4179 (COL). Festuca in South American Paramos Brescia & Marches 4212 (K, P); Farroupilha, in araucarieto, 700 m, 15 Feb 1957, Camargo 59992 (B); Vila Oliva, Caxias do Sul, 24 Feb 1954, Rambo 54999 (B, BAA); Rambo 54989 (B, US); Mun. Cruz Alta, campos de Cruz Alta, 500 m, Feb 1906, Jurgens 6258 (W, US); Serra de Caparas, Espirito Santo, 27Nov 1929, A Chase 10101 (W, US); Sao Leopoldo-Boni Jesus, Dutra 331 (ICN, R); Dutra 402 (R, US); Taimbe, Sao Francisco de Paula, 18 Dec 1950, Rambo 49315 (BAA); Sao Francisco de Paula-Tainhas, Dec 1953, Barreto 1871 (BAA); Mun. de Caxias do Sul, Ana Rech, 780 m, 18 Mar 2000, Scur 656 (US); Taimbesinho, in araucarieto aperto, 20 Feb 1953, Rambo 54011 (US); Esmeralda, estaccao ecologica De Aracuri, 2 Dec 1979, Winge et al. 1362 (ICN); Winge 1272 (ICN); km 60 W of Passo Fundo along highway BR 285 to Vacaria, intersection with Rio Igeiro, 11 Mar 1976, G. Davidse & D 'Arcy 11166 (SP); Sao Francisco da Paula to Eletra, Araucaria woodland 31 Jan 1965, -80 -60 -70 / 1 • *"*%A yJ^*/^^^^ ~*\ ^* 4 " 35 Clayton 4491 (K, SP); Cambara do Sul, 29°00'S, 50°00'W, 27 Jan 1948, Rambo 36446 (K); Cambara do Sul, Itaimbezinho, Bela Vista, 1 Dec 1981, Wagner 944 (ICN); Itaimbezinho, Bela Vista, Vails 2386 (ICN); Wagner etal. 298 (ICN); 1 Dec 1981, Wagner 946 (ICN); Wagner 947 (ICN); Vails 56 (ICN); Lagoa dos Patos, Saco de Tapes, Dec 1980, Goergem & Wagner 50197 (ICN); Mun. de Caxias do Sol-Ana Rech 780 m, 18 Mar 2000, Scur 656 (MO). Santa Catarina: Campo Ere, 29 Feb 1964, Castellanos 24729 (COL, K); Cacador-Curitibanos, 33 km SE of Cacador on the road to Lebon Regis (47 km), 700-900 m, 16 Mar 1957, Smith & Klein 12188 (R, US); Serra de Boa Vista, Sao Jose, 3 Mar 1961, Reitz & Klein 10832 (B, US); 10 Nov 1960, Reitz & Klein 10399 (B, US); Palmares, Campos Novos, 900 m, 11 Apr 1963, Reitz & Klein 14611 (B, US); Mun. Cacador, km W of Cacador, ruderal, 900-1000 m, 6 Feb 1957, Smith & Klein 10887 (R); Bog. 8 km N of Cacador, 950-1100 m, 7 Feb 1957, Smith -50 -40 -30 1 * 10 10 ^^TV^V^J PANAMA \ • VENEZUELA-*; J COLOMBIA^-? Lei 1 <" AJ> f|^v p I) 1 ^iECIMDOR •'y -10 -10 \ PERU \^ ( \A \_ !j BRAZIL -20 -20 m / u ] -30 ^^^^^^ -80 f N 0 ^^^^^^^B ^^^^^^ 1 -70 -60 -50 Figure 11. Distribution of Festuca ulochaeta (•), F. tovariensis (A), and F. caldasii (•). -r 600 Kilometers 4 -40 -30 -30 36 Festuca in South American Paramos & Klein 10956 (R, US); Mun. Joacaba, campos of palmas, 55 km Wof Ca9ador, 1000-1200 m, 18 Feb 1957, Smith & Klein 11384 (R, US); Mun. Chapeco, Fazenda Campo Sao Vicente, 24 km W of Campo Ere, 900-1000 m, 20 Feb 1957, Smith & Klein 11611 (R, US); Mun. Lajes, E of Capao Alto, 900-1000 m, 12 Feb 1957, Smith & Klein 11335 (R, US); Mun. Campo Alegre, lower fazenda of ernesto Scheide, 900 m, 10 Mar 1957, Smith & Klein 12024 (R, US); Morro Juco Prudente, steep slopes of wooded arroyo, 1 Jan 1946, J.R. Swollen 8033 (US); Coxilha Rica, open woods, 6 Jan 1946, J.R. Swollen 8168 (US); Picadas, km 181 da ERF, Papanduva, 1000 m, 26 Feb 1962, Reitz & Klein 12509 (US); Sao Joaquim, Urupema, matinha, 1200 m, 24 Dec 1962, Reitz & Klein 14585 (US); Rio Cacador, 21 Jan 1946, J.R. Swollen 8231 (MO, NY, US); km 21, NE of Santa Cecilia, along Highway BR-116 to Curitiba, open grassland with rocks outcrops, marshy area and patches of trees, 1250 m, 3 Oct 1976, G. Davidse et al. 11082 (NY, SP); km 18, NE of Santa Cecilia, along Highway BR-116 to Curitiba, grassy roadside, 1200 m, 3 Oct 1976, G. Davidse et al. 11087 (NY, SP); Campo Dos Padres, Bom Retiro, 1900 m, 20 Dec 1953, Reitz 2611 (NY). Sao Paulo: 1816-1821, Saint-Hilaire 312 (P); Saint-Hilaire 304 (P); Sao Paulo prope Apiahy, Puiggari s.n. (W); Serra de Bocaina, Apr 1951, Segadas 2827 (R); Campos do Jordao, subida para o pico do Itapeva, Kuhlmann 2242 (SP). COLOMBIA. Cundinamarca: Paramo del Tablazo, W of Subachoque, 3100 m, Wood 3842 (COL, FMB, K); between La Calera- Choachi, ca. 1 km above MundoNuevo, 2600 m, 1983, Wood5104 (COL, K). sin. loc, Lin dig 1862 (P). COSTA RICA. Limon: Canton de Talamanca, Sabanas de Durika, 1 km aguas abajo de la confluencia de los Rio Uk y Rio Kuk, 83°19'30"W, 09°25'20"N, 2250 m, 20 Oct 1989, Herrera 3730 (K, MO). VENEZUELA. Aragua: Colonia Tovar, trayecto El Lagunazo - Colonia Tovar, 2100 m, Feb 1953, Aristeguieta 763 (VEN). Merida: Mun. Tabay, Laguna Coromoto, mountain forest, 3000-3100 m, 7 Nov 2000, D. Stancik 4177 (AAU, CAR, COL, PRC, W); 2800-3000 m, D. Stancik 4179 (CAR, COL, PRC, US); Laguna La Coromoto, 3200-3300 m, 3 Jul 1987, B. Briceno & Adamo 2011 (Herbarium Briceno). 9. Festuca caldasii (Kunth) Kunth, Revis. Gramin. 1:132.1835.(Figs. 11,12,76E&F,77A&B). Bromus caldasii Kunth Nov. Gen. Sp. (quarto ed.) 1:151.1816. Schedonorus caldasii (Kunth) Roem. & Schult, Syst. Veg. 2: 709. 1819. Festuca quadridentata var. caldasii (Kunth) St.-Yves, Candollea 3: 266. 1927. Festuca quitensis Willd. ex Kunth, Enum. PI. 1: 407. 1833, nom. inval. TYPE: Ecuador, crescit locis altis regni Quitensis, prope Chillo, Humboldt & Bonplands.n. (lectotype: P!, designated here; isolectotypes: B!, US-865519 fragm. ex P!). Bromusprocerus Kunth, Nov. Gen. Sp. (quarto ed.) 1: 150. 1816. Schedonorus procerus (Kunth) Roem. & Schult., Syst. Veg. 2:708. 1819. Bromus procerus Humb. ex Spreng., Syst. Veg. 1:357. 1825. nom. illegit. Festuca quadridentata subsp. eminens (Kunth) St.Yves, Candollea 3:266.1927. Festuca eminens Kunth, Revis. Gramin. 1: 132. 1829. TYPE: Ecuador. Pichincha, 2410 m, Feb, Humboldt & Bonpland 2296 (holotype: P!; isotypes: B!, BAA-1206 fragm. ex B, P!, US-2875405 fragm. exP!). Rhizomatous perennials with extravaginal innovations. Culms 70-150 cm tall, erect, scabrous; nodes 3 or 4 in distal half. Leaf sheaths coriaceous, scabrous, chestnut-brown to brown, striate, fibrous near base, margins free; ligules 2.5-3 mm long, membranous, abaxially hairy, apex acute, lacerate; blades 20-35 x 0.3-0.9 cm, flat, green, scabrous, with prickles on both abaxial and adaxial epidermis. Panicles 13-17 x 5-8 cm, ovate; branches erect or sometimes spreading, scabrous. Spikelets 15-17 mm long, oval, florets 5-7(-8); rachilla pilose; glumes 3.5-6.5 mm long, membranous, lanceolate, green with transparent membranous margins; lower glumes 3.5•4(-5) mm long, 1-nerved; upper glumes 4.5-6.5 mm long, 3-nerved; lemmas 10-14 mm long, lanceolate, membranous to coriaceous, 5-nerved, green, scabrous or shortly densely hairy, apex entire, awned, the awn 1-3 mm long; callus glabrous; paleas 4/5 as long as the lemma, membranous, scabrous; lodicules 0.8-1 mm long, oblong-lanceolate; anthers (2.8-)3.5-4.5 mm long; ovary apex glabrous. Caryopses not seen. Leaf blade anatomy.•Cross-sections with many vascular bundles, with just small ribs; sclerenchyma under both abaxial and adaxial epidermis, discontinuous; bulliform cells present, few; epidermis without hairs. Observations.•Festuca caldasii is morphologically similar to F. woodii, a species that occurs in northern Colombia. However, F. woodii has Festuca in South American Paramos 37 1 mm Figure 12. Festuca caldasii. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Laegaard 20405 (F). 38 Festuca in South American Paramos smaller spikelets (12-14 versus 15-17 mm long), shorter lemmas (8.5-9 versus 10-14 mm long), and shorter lemma awns (1 versus 1-3 mm long). In Costa Rica and Panama there are two other species in sect. Glabricarpae: F. breviglumis Swallen and F. chiriquensis Swallen. However, both of these species have shorter (0.5-1 mm long) and acute ligules. Distribution and habitat.•Festuca caldasii occurs sporadically from southern Colombia (Cauca, Narino) to southern Ecuador (Canar, Chimborazo, Loja, Pichincha). This species is known from the margins of Andean forests and matorral vegetation between 1900-3000 m. Additional specimens examined. COLOMBIA. Cauca: Tierra Dentro, below Pitaio, 2400 m, Feb 1906, Pittier 437 (US); Rio Paez Valley, between Huila and Bitonco, 1900-2300 m, Feb 1906, Pittier 1320 (US); near Jambalo, 2200 m, Feb 1906, Pittier 1444 (US). Narino: Mun. Buesaco, a 2 km de la poblacion, 2000 m, Ramirez 1404 (COL); Mun. Pasto, Morasurco hill, 2550 m, 24 Feb 1986, Wood 5309 (COL, FMB, K). ECUADOR. Canar: El Tambo-Carshao road, km 1.5-3, 02°28'S, 78°59'W, 3100 m, 10 Jun 1999, S. Laegaard & Sklenaf 20302 (LOJA); Chimborazo: Canyon of Rio Chanohan, about 5 km N of Hiugra, 5000-6500 ft, CAMP (F, K, NY, US); road Sipambe-Hiugra, km 10, 02°15'S, 78°57'W, 2050 m, S. Laegaard 20405 (AAU, PRC); Huigra, 1200 m, A.S. Hitchcock 20746 (US); km 6 NE of Pallatanga and 5.4 km W on road to Chillanes, 1950 m, P.M. Peterson & E.J. Judziewicz 9259 (QCA, US). Canon of the Rio Chanchan, about 5 km N of Huigra, moist forested valleys in the afternoon fog belt, 5000-6500 ft, 19-28 May 1945, Camp 3327 (K). Loja: Road Catacocha-La Toma, km 28,03°58'S, 79°31 'W, 2200-2250 m, S. Laegaard 102535 (AAU, QCA); Zumba road, km 6 above Jimbura, 04°40', 79°26'W, 2400-2450 m, S. Laegaard 105256 (AAU); road Amalusa-Jimbura, km 7-9, 04°36'S, 79°28'W, 1900 m, S. Laegaard 105235 (AAU, QCA). Pichincha: Cotocollao, Sodiro s.n. (QPLS, US); Sodiro s.n. (W). 10. Festuca reclinata Swallen, Contr. U.S. Natl. Herb. 29(6): 254.1949. (Figs. 13,14,77C-F). TYPE: Colombia. Santander: Paramo de Almorzadero, Cordillera Oriental, 3500-3700 m, 20 Jun 1940, J. Cuatrecasas & KG. Barriga 9970 (holotype: US-1798714!; isotype: COL!). Rhizomatous perennials forming small tussocks with extravaginal innovations. Culms 30^10 cm tall, decumbens to erect, glabrous; nodes 2 or 3 in distal half; leaf sheaths membranous, greenish-white, scabrous, upper sheaths closed for 1/2 the length; auricles absent; ligules 1-2.5 mm long, membranous, apex acute, ephemeral; blades 5-15 x 0.3-0.5 cm, flat, green, abaxially scabrous. Panicles 9-10 cm x 2-3 cm, flexuous, ovate, branched; branches glabrous. Spikelets 10-13 mm long, obovate, florets 4; rachilla papillose; glumes 1.3-3.5 mm long, membranous, lanceolate, green, glabrous, upper margins hairy; lower glumes 1.3-2 mm long, 1 -nerved; upper glumes 3-3.5 mm long, 3-nerved; lemmas 7-8.5 mm long, 5-nerved, lanceolate, membranous, green, scabrous, apex two-dentate, awned between the teeth, the awn 1-2 mm long; callus glabrous; paleas 4/5 as long as the lemma, membranous, keels scabrous; lodicules lanceolate; anthers 3-3.5 mm long; ovary apex glabrous. Caryopses not seen. Leaf blade anatomy.•Cross-sections with many vascular bundles, with small ribs; sclerenchyma under both abaxial and adaxial epidermis, discontinuous, small, extending to the vascular bundles forming girders; bulliform cells absent; epidermis sparsely hairy. Observations.•Festuca reclinata superficially resembles Aphanelytrum procumbens Hack. The spikelets and panicles of F. reclinata are similar to Aphanelytrum, but the glumes in the former are clearly nerved (versus unnerved), which is the difference between the two genera. Alexeev (1986) placed F. reclinata in sect. Glabricarpae and we consider the Alexeev's decision to be only provisional. Distribution and habitat.•Festuca reclinata is known only from the type locality in the Colombian Cordillera Oriental, Dept. Santander were it was found in a paramo. 11. Festuca woodii Stancik, Darwiniana 41(1•4): 107. 2003. (Figs. 14, 15, 78A-D). TYPE: Colombia. Boyaca, Sierra Nevada del Cocuy, Hda. La Esperanza, 3700 m, in crevices of limestone pavement on a steep, open dip slope. Vigorously tufted perennial more than 1 m, inflorescence purple brown, 29 Oct 1985, J.R.I. Wood 5254 (holotype: COL!, isotypes: FMB!,K!). Rhizomatous perennials with extravaginal innovations. Culm 70-90 cm tall, erect, scabrous; Festuca in South American Paramos 39 1 mm Figure 13. Festuca reclinata. A. Growing form. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf blade cross-section. A-G, Cuatrecasas & Barriga 9970 (COL). Festuca in South American Paramos 40 -85 -80 -75 -70 Figure 14. Distribution of Festucapratensis (•), F. woodii (A), and F. reclinata (•). nodes 2 or 3; leaf sheaths membranous, glabrous, grayish, fibrous and ephemeral at base, margins free; auricles absent; ligules ca. 0.5 mm long, membranous, apex truncate; blades 50-60 x 0.25-0.5 cm, flat, conduplicate near apex, green, scabrous, with prickles on both abaxial and adaxial surface. Panicles ca. 20 cm long and 15 cm wide, flexuous, ovate, branched; branches scabrous. Spikelets 1214 mm long, ovate, florets 4 or 5; rachilla densely hairy; glumes 4.5-7.5 mm long, membranous, narrowly lanceolate, green, sparsely scabrous; lower glumes 4.5-5.5 mm long, 1-nerved; upper glumes 6-7.5 mm long, 3-nerved; lemmas 8.5-9 mm long, 5-nerved, membranous to coriaceous, lanceolate, green, scabrous and short-hairy, apex two-dentate, awned, the awn 0.5-1 mm long; callus glabrous; paleas as long as the lemma, membranous, papillose; anthers 3.5-4 mm long; ovary apex sparsely short-hairy. Caryopses not seen. Leaf blade anatomy.•Cross-sections with many vascular bundles and small ribs above; sclerenchyma under both abaxial and adaxial epidermis, discontinuous, extending to the vascular bundles forming girders; bulliform cells present; epidermis without hairs. Observations.•Festuca woodii is morphologically similar to F. caldasii known from southern Colombia and Ecuador. However, F woodii has shorter ligules (0.5 versus 2.5-3 mm long), truncate (versus acute) ligules, smaller spikelets (12-14 versus 15-17 mm long), and longer glumes. Distribution and habitat.•Festuca woodii is endemic to the Colombian Cordillera Oriental (Boyaca, Cundinamarca) occuring in matorral and grass paramo vegetation types on rocky slopes and calcareous outcrops between 2700-3700 m. Additional specimens examined. COLOMBIA. Cundinamarca: Bogota, 2730 m, 16 Sep 1915, Festuca in South American Paramos 41 Figure 15. Festuca woodii. A. Stylized growth form. B. Habit. C. Spikelet. D. Glumes. E. Lemma. F. Leaf blade cross-section. A-F, Wood 5254 (COL). 42 Festuca in South American Paramos Apollinaire & Arthur 18 (US). Boyaca: Sierra Nevada del Cocuy, Hda. La Esperanza, 3700 m, 29 Dec 1985, Wood 5354 (FMB). 12. Festuca arundinacea Schreb., Spic. Fl. Lips. 57. 1771. (Figs. 16,17, 78E & F, 79A & B). Bromus arundinaceus (Schreb.) Roth, Tent. Fl. Germ. 2: 141. 1789. Schedonorus arundinaceus (Schreb.) Dumort., Observ. Gramin. Belg. 106.1824, nom. conserv. Festuca elatior var. arundinacea (Schreb.) Wimm., Fl. Schles. 3: 59. 1857. Festuca elatior subsp. arundinacea (Schreb.) Celak., Prodr. Fl. Bohmen 1: 51. 1867. Festuca elatior subsp. arundinacea (Schreb.) Hack., Monogr. Festuc. Eur. 152. 1882. Lolium arundinaceum (Schreb.) Darbysh, Novon 3(3): 241.1993. TYPE: Scheuzer, Agrostographia, tab. 5, fig. 18. 1719. (lectotype: designated by Reveal, Terrell, Wiersema & Scholz, Taxon40: 136. 1991). Festuca elatior L., Sp. PI. 1:75.1753, nom. rej. Poa elatior (L.) Moench, Enum. PI. Hess. 37.1777. Avena secunda Salisb., Prodr. Stirp. 22. 1796. Bromus elatior (L.) Koeler, Descr. Gram. 214. 1802. Schedonorus elatior (L.) P. Beauv., Ess. Agrost. 99, 156, 162, 177. 1812. Festucapratensis var. elatior (L.) Gaudin, Fl. Helv. 1:293. 1828. Bucetum elatius (L.) Parnell, Grasses Scotl. 107, pi. 46. 1842. Tragus elatior (L.) Panz. ex B.D. Jacks., Ind. Kew. 2: 1098. 1895. Gnomonia elatior (L.) Lunell, Amer. Midi. Nat. 4: 224. 1915. (lectotype: LINN-92.17, designated by Terrell, Brittonia 19: 131. 1967; again by Linder, Bothalia 16: 59. 1986). Poa phoenix Scop., Fl. Carniol., ed. 2,1: 74. 1771. Schedonorus phoenix (Scop.) J. Holub, Preslia 70: 113. 1998. Bromus littoreus Retz., Fl. Scand. Prodr. 19. 1779. Schedonorus littoreus (Retz.) Tzvelev, Nov. Sist. Vyssh. Rast. 31: 258. 1998. Festuca littoralis Wahlenb., nom. illeg. horn., Nova ActaRegiae Soc. Sci. Upsal. 8: 211. 1821. Festuca pseudosclerophylla Krivot., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk SSSR 17: 73. 1955. Leucopoa pseudosclerophylla (Krivot.) Bor in K. H. Rech, Fl. Iranica 70: 73. 1970. Poa hybrida var. vallesiaca Bronm., Repert. Spec. Nov. Regni Veg. 16: 301. 1919. Festuca elatior subsp. arundinacea var. genuina subvar. orientalis Hack., Monogr. Festuc. Europ. 154. 1882. Festuca orientalis (Hack.) Krecz. & Bobrov, Fl. URSS 2: 531. 1934. Festuca arundinacea subsp. orientalis (Hack.) Tzvelev, Fl. URSS 18: 17. 1970. Festuca regeliana Pavl., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 41(1): 80. 1938. Loosely tufted to shortly rhizomatous perennials with extravaginal innovations. Culms (50-) 100-150 cm tall, erect, scabrous; nodes 3. Leaf sheaths coriaceous, striate, glabrous or scabrous; auricles present, falcate, margins ciliate; ligules ca. 0.5 mm long, membranous to coriaceous, apex truncate; blades 25-30 cm long, 5-11 mm wide, flat, green, abaxially scabrous. Panicles 10-17 x 4-6 cm, narrow, with erect branches; branches scabrous. Spikelets 9-12(-15) mm long, florets 6-8(-10); rachilla scabrous; glumes 3.3-6.5(-7)mm long, lanceolate, membranous to coriaceous, green, glabrous, apex acute; lower glumes 3.3•5(•5.5) mm long, 1-nerved; upper glumes 4.5-6.5 mm long, 3-nerved, sometimes scabrous on back; lemmas 6-7.5(-8) mm long, 5-nerved, lanceolate, membranous to coriaceous, green, apex scabrous on midrib, mucronate or short-awned, the awn 0.5-1.5 mm long; paleas almost as long as the lemma, scabrous on margins and keels; lodicules lanceolate, acuminate; anthers 2.7-3.5 mm long; ovary apex glabrous or sparsely hairy. Leaf blade anatomy.•Cross-sections with numerous vascular bundles, with small ribs above; sclerenchyma discontinuous, extending to the vascular bundles under both abaxial and adaxial epidermis; bulliform cells present. Observations.•This species is often treated by many authors as a separate genus, Schedonorus arundinaceus (Soreng et al. 2003). Distribution and habitat.•Festuca arundinacea is introduced from Europe and cultivated in pastures, fields and rarely escaping along roadsides between 2300-3600 m. Additional specimens examined. COLOMBIA. Cundinamarca: Mun. de Suba, Hda. Las Mercedes, 16 May 1964, E. Forero et al. 45 (COL); 8 Oct 1964, E. Forero et al. 82 (COL); 15 Oct 1964, E. Forero et al. 92 (COL); between Guasca and La Calera, Vereda Santa Helena, 2950 m, 9 Dec 1984, Wood 4639 (COL, K). Narifio: Mun. Ipiales, "Las Lajas", 2740 m, 8 Aug 1939, Garcia-Barriga 7838A (COL). ECUADOR. Cafiar: S of El Tambo, km 1.5-3 on road to Carshao, 02°28 'S, 78°59'W, 3100 m, S. Laegaard & Sklendr 20301 (AAU); Mun. El Tambo, road from El Tambo to Ingapirca, km 1, Festuca in South American Paramos 43 2 mm Figure 16. Festuca arundinacea. A. Stylized growth form. B. Habit. C. Inflorescence. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 4102 (PRC). 44 Festuca in South American Paramos burned matorral and paramo, 3400-3500 m, 29 Aug 2000, D. Stancik 3798 (PRC, QCA). Chimborazo: km 5 N of Tixan, meadows, 02°06'S, 78°46'W, 3280 m, S. Laegaard 101818 (AAU, QCAQCNE); S. Laegaardetall03003 (AAU, QCA, QCNE); km 22 on road Alausi-Riobamba, 02°08'S, 78°46'W, 3250 m, S Laegaard 18667 (AAU,LOJA, QCA, QCNE); pasture near Riobamba, Sodiro s. n. (QPLS). Cotopaxi: Salcedo, 2650 m,Acosta-Solis 10206 (V, US). Loja: La Argelia (Escuela de Agronomia), 04°02'S, 79°12'W, 2100 m, S. Laegaard 18704 (LOJA). Morona-Santiago: Hda. Huargualla-Hda. SanEduardo, 01°57'S, 78°32.2'W, 3600 m, 19 Jul 1999, D. Stancik 3309 (PRC, QCA); D. Stancik 3310 (PRC, QCA). Pichincha: Quito, 00°10'S, 78°30'W, 2850 m, 30 Mar 1998, S. Laegaard 18623 (LOJA); Quito-Santa Catalina, 00°22 'S, 78°21 'W, Feb 1981, Vivar & Marin 1305 (LOJA); Quito-El Batan, 2850 m, Acosta-Solis 19821 (US); AcostaSolis 19832 (US); Acosta-Solis 19835 (US); W side of Mt. Pichincha, 3070 m, Sodiro s.n. (QPLS); Pichincha, Aug ISSS, Sodiro s.n. (QPLS); 3000 m, 1904, Sodiro s.n. (US); Road Lloa-Guagua Pichincha, km 6,00°13 'S, 78°35 'W, 3600 m, S Laegaard 102716 (AAU). Mun. Otavalo, Ruchanda-road to Quito, 3100m, 14 Jul 1999..D. Stancik3221 (PRC, QCA); Mun. Otavalo, road from Laguna Mojanda to Cochasqui, 00°04'55"N, 78°17'50"W, 3450 m, 19 Oct 2000, D. Stancik 4102 (PRC, QCA). Tungurahua: Tungurahua, Aug 1901, Sodiro s.n. (QPLS, US); Slope of Mt. Tungurahua above Banos, 2300 m, E. Asplund8425 (S); Pillaro, 2850 m, E. Asplund 8150 (NY, QCA, S). Zamora-Chinchipe: Old road Loja-Zamora ca. 1 km E of pass, 03°59'S, 79°09'W, 2750 m, 26 Apr 1998, S Laegaard 18744 (AAU, LOJA, QCA, QCNE). VENEZUELA. Barinas: 74 km NW of Barinas on Hwy 1 and 71 km NE of Merida, slopes with Festuca, Espeletia, Loutegia, Stevia, and Sporobolus, 2740 m, 24Nov 1991, P.M. Peterson 11181 (US). Merida. Entre Pedregal y Apartaderos, praderas de fuentes del Chama, asociado aJuncus, Rumex etc., 3240 m, 23 Aug 1981, Ponce & Trujuillo 245 (MY); Dept. Rangel, Paramo de Mucubaji, Mesa del Caballo, El Pedregal, 3350 m, 12 Jun 1981, B. Briceno & Adamo 286 (Herbarium Briceno, PRC). 13. Festuca pratensis Huds., Fl. Angl. 37. 1762. (Fig. 14). Festucafluitans var.pratensis (Huds.) Huds., Fl. Angl. (ed. 2) 47. 1778. Schedonorus pratensis (Huds.) Beauv., Ess. Agrostogr. 99, 163, 177. 1812. Bromus pratensis (Huds.) Spreng., Syst. Veg. 1: 359. 1825, nom. illeg. Bucetum pratense (Huds.) Parnell, Grass. Scotland 105, t. 46. 1842. Festuca elatior var. pratensis (Huds.) A. Gray, Man. Bot. (ed. 5) 634. 1867. Festuca elatior subsp. pratensis (Huds.) Hack, Bot. Centralbl. 8: 407. 1881. Festuca elatior subsp.pratensis (Huds.) Hack, Monogr. Festuc. Eur. 150.1882,isonym. Traguspratensis (Huds.) Panz. ex B.D. Jacks, Index Kew. 2:1099. 1895. Loliumpratense (Huds.) Darbysh, Novon 3(3): 242. 1993. TYPE: Great Britain (BM-SL), Buddie s.n., (lectotype: BM-SL!, Herb. Sloane. 125.16, designated by Reveal, Terrell, Wiersema &Scholz,Taxon40:135. 1991). Festuca poaeoides Michx, Fl. Bor.-Amer. 1: 67. 1803, nom. illeg. Festuca poaeoides var. americana Pers, Syn. PI. 1:94.1805. Festuca americana (Pers.) F G. Dietr, Nachtr. vollst. Lex. Gartn. 3: 332.1817. Schedonorus americanus (Pers.) Roem. & Schult, Syst. Veg. 2: 706. 1817. Schedonorus radicans Dumort, Obs. Gramin. belg.: 106. 1824. Festuca radicans (Dumort.) Steud, Syn. PI. Glum. 1: 309. 1854. Festuca glabra Spreng, Syst. Veg. 1: 353. 1824, nom. illeg. horn. Rhizomatous perennials with extravaginal innovations. Culms 60-120 cm tall, erect, scabrous; nodes 3 in the basal half. Leaf sheaths (lowermost) coriaceous, brown, striate, scabrous, margins free, sometimes fibrous; auricles falcate, without hairs on margins; ligules 0.3-0.5 mm long membranous, sometimes coriaceous, apex truncate; blades 20-25 x 0.4-0.8(-l) cm, flat, green, abaxially scabrous. Panicles 15-20 x 3-5 cm, contracted, narrow; branches scabrous. Spikelets 9-11 mm long, florets 5 or 6; rachilla scabrous; glumes 2.7-4 mm long, lanceolate, membranous, sometimes coriaceous, green, glabrous, apex acuminate; lower glumes 2.7-3 mm long, 1-nerved; upper glumes 3.7-4 mm long, 3-nerved; lemmas 6-6.5 mm long, 5-nerved, membranous to coriaceous, lanceolate, green, apex scabrous, mucronate; paleas almost as long as the lemma, scabrous on margins and keels; lodicules lanceolate, acuminate; anthers 2.7-3.5 mm long; ovary apex glabrous or sparsely hairy. Leaf blade anatomy.•Cross-sections with numerous vascular bundles, without ribs above; sclerenchyma under both abaxial and adaxial epidermis, discontinuous, extending to the vascular bundles forming girders; bulliform cells present. Festuca in South American Paramos -85 -80 -75 -60 -65 -70 -55 & } ^ZJ^K'*' •A PANAMA 45 10 10 /? • lN*\j (m VENEZUELA COLOMBIA i\. *f ECUADC BRAZIL I PERU % V,u^- -10 N 0 300 600 Kilometer \ -< k - -85 -80 -75 -70 f -10 -65 a -60 -55 Figure 17. Distribution of Festuca dichoclada (•), F. arundinacea (A), and F. venezuelana (•). Observations.•Modern authors generally treat this taxon as a separate genus, Schedonorus pratensis (Soreng et al. 2003). Distribution and habitat.•This species is introduced from the Europe and sporadically cultivated at an altitude around 3000 m. Additional specimens examined. ECUADOR. Chimborazo: Riobamba, Sodiro s.n. (US). Pichincha: Quito-Santa Catalina, 00°22'S, 78°31'W, Vivar & Marin 1306 (LOJA); Canton TabacundoPicalqui, 2750 m, Acosta-Solis 16268 (US). 14. Festuca dichoclada Pilg., Bot. Jahrb. Syst. 37: 514. 1906. (Fig. 17). Festuca quadridentata var. dichoclada (Pilg.) St.-Yves, Candollea 3: 265. 1927. TYPE: Peru. Ancachs (Ancash), in declivibus montium Cordillera blanca supra Caraz in faucibus umbrosis fruticibus altis obtectis, 3300-3600 m, 9 Jun 1903, A. Weberbauer 3230 (holotype: B!; isotypes: BAA-1196 fragm. ex B, US-2875396 fragm. exB!). Large rhizomatous, loosely caespitose perennials with intravaginal innovations. Culms 150-200 cm tall, erect, scabrous; nodes 3 or 4 in distal half. Leaf sheaths coriaceous, scabrous; auricles absent; ligules 8-12 mm long, membranous, apex acute; blades 40-100 x 0.5-1.4 cm, flat occasionally partially conduplicate, green, abaxially scabrous, with prickles on abaxial and adaxial epidermis. Panicles 20-40 x 20-25 cm, ovate; branches scabrous. Spikelets 11-14 mm long, oblong, florets 3-5; rachilla hairy; glumes 5-9 mm long, lanceolate, greenishwhite, smooth to papillose, apex acute; lower glumes 5-7.5 mm long, 1-nerved; upper glumes 6.5-9 mm long, 3-nerved; lemmas 8.5-10 mm long, lanceolate, 5-nerved, membranous to coriaceous, scabrous, apex entire, awnless; callus glabrous; paleas as long as the lemma, smooth to papillose; 46 Festuca in South American Paramos lodicules ovate, obtuse; anthers 3.5^.2(-4.7) mm long; ovary apex glabrous. Caryopses oblong; hilum nearly as long as the caryopsis. Leaf blade anatomy.•Cross-sections with many vascular bundles and ribs above; sclerenchyma under both abaxial and adaxial epidermis, discontinuous; bulliform cells present; epidermis without hairs. Distribution and habitat.•Known only from northern and central Peru. It occurs along margins of Andean forests, clearings, rocky slopes, and paramos between 3000-4000 m. Additional specimens examined. PERU. Ancash: Prov. Bolognesi, Jerusalem, cerro al E de Aquia, arcilloso-pedregoso en monte pluvifolio, 3200-3250 m, 18 May 1950, R. Ferreyra 7526 (US); Casca, abajo de Chiquian, 3100-3200 m, 9 May 1950, R. Ferreyra 7281 (US); Prov. Yungay, Huascaran Parque Nacional, between Lake Llanganuco and Portachuelo, 77°36'W, 09°03'S, 4200-4329m, 16 Aug 1984, Smith 8233 (US). Prov. Huaraz, Huascaran Parque Nacional, Quebrada Shallap, 77°24'W, 09°30'S, 3700^000 m, 22 May 1985, Smith et al. 10769 (F); Prov. Huaylas, Huascaran Parque Nacional, environs of Auquispuquio, 77°57'W, 08°49'S, 3900^000 m, 4 Sep 1986, Smith et al. 12110 (F); Prov. Yungay, Huascaran Parque Nacional, Llanganuco sector, Maria Josefa trail between Chinancocha and Pucayacu, 77°39'W, 09°05'S, 3700-3850 m, 5 Jul 1985, Smith 10537 (F); Prov. Carhuaz, Huascaran Parque Nacional, Quebrada Ulta, on road to Ulta Pass, 77°32'W, 09°07'S, 4000-4400 m, 29 Jul 1985, Smith 11389 (F). Cajamarca: Prov. Contumaza, Guzmango, cerro Campanulas, 3050 m, 1 Jun 1959, SagdsteguiiSagdsteguii 2991 (US); Prov. Cajamarca, SAIS Jose Carlos Mariategui, km 20^t0 on Sunchubamba-San Juan road, jalca with small patches of ceja de selva in rocky area, 3300-3500 m, 5 Jun 1984, Smith 7534 (US); Prov. Cajamarca, Distr. San Juan, carretera San JuanCajamarca, arriba de San Juan, ladera arcillosapedregosa, 2350 m, 6 Dec 1993, Vega 730 (F); Prov. Cajamarca, Cumbe Mayo, 21 km al W de Cajamarca, ladera con arbustos disersos, 3100 m, 4 Nov 1977, Vega 1966 (V), Huancavelica: Conaica, Carhuay-pampa arriba de Conaica, 3700-3750 m, 18 Mar 1951, O. Tovar 143 (US). Junin: Prov. Huancayo, Hda. Acopalca, 4000 m, 20 Jul 1945, Infantes 435 (US). La Libertad: Prov. De Otuzca, cerca a Usquil, falda de cerro, junto con Chusquea, Loasa, Cajophora, 3000-3100 m, 9 Jun 1950, R. Ferreyra 7642 (K, US); R. Ferreyra 7664 (K, US); Otuzco, entre piedras, 2600 m, 6 Apr 1990, Leiva & Leiva 96 (F); Otuzco, cerro de los Enamorados (al norte de Salpo), ladera, 3440 m, 16 Jun 1993, Leiva 797 (MO); Near Usquil, Utusco, hillside, 3200 m, 9 Jun 1950, Anderson 1271 (US). 15. Festuca quadridentata Kunth, Nov. Gen. Sp. (quarto ed.) 1: 154. 1816. (Figs. 18, 21, 79C-F, 80A). Festuca flexuosa Willd. ex Kunth, Enum. PI. 1: 407. 1833, nom. inval. pro syn. TYPE: Ecuador. Chimborazo, 2860 m, Humboldt & Bonpland s.n. (holotype: P!; isotypes: B!, P!). Large tussocked perennials with intravaginal innovations. Culms 170-200 cm tall, erect, scabrous; nodes 3 or 4 in distal half. Leaf sheaths coriaceous, striate, scabrous; auricles absent; ligules (8•)9•13 mm long, membranous, apex acute; blades 40-150 x 0.7-1.4 cm, flat to conduplicate, green, scabrous with prickles on abaxial and adaxial epidermis. Panicles 20•^0 x 15-20 cm, ovate; branches scabrous. Spikelets 11•13(•15) mm long, oblong, florets (3-)4 or 5 (-6); rachilla with short, scattered hairs; glumes (3.5-)4-6.5 mm long, lanceolate, greenish-white, smooth sometimes papillose, apex acute; lower glumes (3.5-)4•5 mm long, 1-nerved; upper glumes 5-6.5 mm long, 3-nerved; lemmas 8.5-9.5 mm long, lanceolate, 5-nerved, membranous to coriaceous, scabrous or papillose, apex erose and dentate, awnless or sometimes mucronate; callus glabrous; paleas as long as the lemma or longer, smooth or papillose; lodicules ca. 0.7 mm long, ovate, apex obtuse; anthers 3.5•4.2(•4.7) mm long; ovary apex glabrous. Caryopses oblong; hilum nearly as long as the caryopsis. Leaf blade anatomy.•Cross-sections with many vascular bundles, corresponding to the number of ribs above; sclerenchyma under both abaxial and adaxial epidermis, discontinuous; bulliform cells present; epidermis without hairs. Observations.•Festuca quadridentata is morphologically similar to the Peruvian species F dichoclada, however the former differs by having shorter lower glumes (5-7.5 mm long in F dichoclada), shorter upper glumes (3.5-5 mm versus 6.5-9 mm), and dentate lemmas (versus entire). Distribution and habitat.•This species is endemic to central Ecuador (Canar, Chimborazo, Morona-Santiago, Pichincha) where is known from Festuca in South American Paramos 47 Figure 18. Festuca quadridentata. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-G, Laegaard & Sklendf 20308 (PRC); H, Stancik 3317 (PRC). 48 Festuca in South American Paramos Andean mountain forests, between (1500-)30003400 m. Additional specimens examined. ECUADOR. Caflar: S of El Tambo, ca. 1.5-3 km road to Carshao, 02°28'S, 78°59'W, 3100 m, 10 Jun 1999, S. Laegaard & Sklendf 20308 (AAU, LOJA, PRC); 10 Jun 1999, S. Laegaard & Sklendf 20287 (AAU, LOJA, PRC). Chimborazo: near Alao, along Rio Alao, 0P52'S, 78°30'W, 3200-3400 m, 22 Sep 1985, S. Laegaard 55290 (AAU, QCNE); km 10 N of Sipambe, 02°08'S, 78°52'W, 3400 m, 11 Nov 1985, S. Laegaard 55567 (AAU, QCA); road Alao-Chambo, km 10, 01°50'S, 78°34'W, 3000 m, 11 Oct 1985, S Laegaard 55411 (AAU, QCA); in prov. Riobamba, Sodiro s.n. (QPLS); Alao, Hab. Llactapamba, 3200 m, Acosta-Solis 7581 (F, US); eastern Cordillera of Riobamba, 3200 m, Rimbach 54 (F, US). Morona-Santiago: Parque Nacional Sangay, Hda. Huargualla-San Eduardo, 02°00.25'S, 78°27'W, 3700 m, D. Stancik 3317 (AAU, QCA, PRC, US). Pichincha: Gualea, Sodiro s.n. (QPLS); 1500 m, Mille s.n. (QPLS). Sin. loc. Sodiro 20/5 (P). glabrous. Caryopses oblong-lanceolate; hilum 3/4 of caryopsis length. Leaf blade anatomy.•Cross-sections with many vascular bundles, with shallow ribs; sclerenchyma under both abaxial and adaxial epidermis, discontinuous; bulliform cells present; epidermis with prickles on abaxial and adaxial surfaces, macrohairs absent. Observations.•Festuca venezuelana is morphologically similar to the Bolivian F. steinbachii. However, F steinbachii has shorter ligules (ca. 1 mm long), shorter spikelets (11-13 mm long), and shorter awns (0.5-1.3 mm long). Distribution and habitat.•Festuca venezuelana ranges from Colombian Cordillera Oriental (N de Santander) to the western part of the Venezuelan Andes (Tachira). It occurs in Andean forest clearings between 2800-3400 m. Additional specimens examined. COLOMBIA. Norte de Santander: between Mutiscua and Pamplona, edge of wood, 3400 m, 32 Feb 1927, Killip 19719 (K, US). VENEZUELA. Tachira: Grita, Porqueras-AldeaAguaDiaz, 22 Sep 1942, Tamayo 2326 (US, VEN). 16. Festuca venezuelana Stancik, Darwiniana 41(1^): 111, f. 15b-l. 2003. (Figs. 17, 19, 80B-F). TYPE: Venezuela. Tachira, Mun. La Grita, Paramo La Negra, cross of the roads to La GritaandPogonero, 08°13'22"N, 7P52'51"W, shrubby margin of the road with Asteraceae, Melastomataceae, Cordia sp., 2800 m, 11 Nov 2000, D. Stancik 4262 (holotype: PRC!, isotypes: AAU!, CAR!, COL!, W!). 17. Festuca fragilis (Luces) B. Briceno, Ernstia 4(3^): 7 8-79. 1994. (Figs. 20, 21, 81A-F). Helleria fragilis Luces, J. Wash. Acad. Sci. 32(6): 157, f. 1.1942, MOOT, inval. Hellerochloa fragilis (Luces) Rauschert, Taxon 31(3): 561. 1982. TYPE: Venezuela. Merida, Paramo de Tucani, 4500 m, 17 Dec 1910, A. Jahn 62 (holotype: VEN!; isotype: US!). Tussocked perennials with extravaginal innovations. Culms 130-180 cm tall, erect, scabrous; nodes 3. Leaf sheaths coriaceous, fibrous, scabrous, brown; auricles absent; ligules 3-5 mm long, membranous, apex acute; blades 25^10 x 0.5-1.1 cm, flat, green, abaxially scabrous. Panicles 20-25 x 10-15 cm, lax, nutant; branches scabrous. Spikelets 14-16 mm long, oblong-lanceolate, florets 5-7; rachillas 1-1.2 mm long, scabrous; glumes 3-9 mm long, narrowly lanceolate, coriaceous, margins membranous, apex acute; lower glumes 3•4.5 mm, 1-nerved; upper glumes 7-9 mm long, 3-nerved, apex scabrous; lemmas 10-11 mm long, 5-nerved, lanceolate, coriaceous, brownish-green, scabrous, apex two-dentate and awned, the awn 2-8 mm long, some plants awnless; paleas as long as the lemma, scabrous on margins and keels; lodicules lanceolate, acuminate; anthers 3-3.5 mm long; ovary apex Small tussocked perennials with intravaginal innovations. Culms 20-25 cm tall, erect, finely scabrous; nodes 1, basal. Leaf sheaths membranous to coriaceous, stramineous, glabrous, inconspicuously striate; auricles absent; ligules 4-5 mm long, membranous, apex acute, ciliate; blades 10-15 cm long, 0.6-0.7 mm wide, conduplicate to involute, abaxially scrabrous, olive-green, only 2 or 3 per culm, apex acute. Panicles 5-8 x 1-2 cm, contracted, dense, lanceolate, branches scabrous. Spikelets ca. 3 cm long, obovate, florets 5-7; rachilla densely hairy; glumes 7.5-13 mm long, membranous, lanceolate, margins dentate, apex acute; lower glumes 7.5-9.5 mm long, 1-nerved; upper glumes 10-13 mm long, 3-nerved; lemmas 17-20 mm long, 5-nerved, lanceolate, membranous, upper 1/2 and margins scabrous, apex shortly two-dentate and awned, the awn 5-7 mm long; callus glabrous; Festuca in South American Paramos 49 fHil? 1 mm Figure 19. Festuca venezuelana. A. Stylized growth form. B. Habit. C. Inflorescence. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 4263 (PRC). 50 Festuca in South American Paramos Figure 20. Festuca fragilis. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf blade cross-section. A-G, from Stancik 4192 (PRC). Festuca in South American Paramos -80 -75 -70 51 -65 Figure 21. Distribution of Festuca quadridentata (•), F.fragilis (A), an&F. rubra (•). paleas 3/4 as long as the lemma, membranous, lanceolate, papillose, upper third and along keels scabrous; lodicules lanceolate; anthers 1.5-2.2 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 2/3 of total length. Leaf blade anatomy.•Cross-sections with 7 vascular bundles and 5 ribs above; sclerenchyma under abaxial and adaxial epidermis, continuous, girders absent; epidermis with hairs on adaxial surface, scattered, the hairs 0.03 mm long. Observations.•The Mexican species, F. livida (F subg. Helleria) is morphologically similar to F.fragilis but differs from it by having shorter culms (10-15 cm tall), shorter lemmas (8-14 mm long), and shorter lemmatal awns (1-4 mm long). Distribution and habitat.•Festuca fragilis is known only from the state of Merida, Venezuela and the Sierra Nevada del Cocuy, Colombia, from dry rocky slopes of super-paramo between 3600^1800 m. Additional specimens examined. COLOMBIA. Arauca: Sierra Nevada del Cocuy, Laguna La Plaza, 4300 m, 31 Dec 1985, Wood 5259 (COL). VENEZUELA. Merida: Mun. Mucuchies, Paramo de Piedras Blancas, mountain ridge towards Alto Mucunano, 08°49'54"N, 70°57'33"W, 4350 m, 4 Nov 2000, D. Stancik 4182 (CAR, COL, PRC, US, W); Laguna Negra, 08°49'16"N, 70°57'02"W, 4270 m, D. Stancik 4192 (AAU, CAR, COL, PRC); Mun. Merida, Sierra Nevada-Pico Espejo, 8°42'N, 71°12'W, 4 Nov 2000, D. Stancik 4248 (AAU, CAR, COL, PRC, W); Barclay & Juajibioy 10180 (US); Pico de Las Piedras Blancas, 4740 m, 25 May 1952, Vareschi 1232 (VEN); Camino a Pico Bolivar, 4100-4300 m, 10 Oct 1953, Little 15717 (MER); paramo Media Luna, pendiente N del Pico 52 Festuca in South American Paramos El Toro, 08°32'27"N, 71°04'19"W, super-paramo con Lachemilla equisetifolia y Coespeletia moritziana, 4350 m, 21 Dec 1994, Berg 470 (K); Sierra Nevada de Santo Domingo, between Apartaderos and Timotes, E facing rocky slope, 21-26 Nov 1959, Barclay et al. 9677 (US); Laguna Coromoto, 3600 m, Oct 1956, Aristeguieta 2596 (VEN); 3400 m, Oct 1956, Aristeguieta 2603 (VEN); Paramos mas altos cerca de El Gavilan, 4200 m, 25 Jan 1929, Pittier 13276 (VEN, US); Paramo de Piedras Blancas, 4300 m, 25 Sep 1952, Vareschi et al. 1233 (VEN); 4200 m, 28 Feb 1976, Baruch 46 (VEN); 4400 m, 8 Dec 1979, Barreto 656 (MERC); 4000^600 m, 31 Oct 1981, B. Briceno et al. 358 (MERC); 4150^250 m, 13 Nov 1976, Ricardi & Carrez 5930 (MER); Paramo de Mucuchies, Pico Aguila, Pico de Aguila, 3800-3900 m, 22 Jan 1986, Bono 5662 (VEN); 4300 m, 17 Oct 1984, Bono 4328 (VEN); 4700 m, 12 Feb 1976, Teran 13215 (MERF); 4118m, 6 Dec 1980, Badillo et al. 7562 (MY); Pico Espejo, 4765m, 17 Dec 1969,/. Steyermark & Koyama 102389 (VEN); 4680^1780 m, 31 Dec 1961, Teran & Melchior 833 (MER, HERZ); Teran & Melchior 848 (MER); 4400 m, 16 Dec 1952, Bernardi 268 (MER); Loma Redonda-Alto de La Cruz, 4040-4300 m, 11 Nov 1994, B. Briceno 3074 (Herbarium Briceno); Mun. Rangel, Paramo El Banco, entrada por la Toma baja, 4420 m, 21 Oct 1997, B. Briceno & Molinillo 3461 (Herbarium Briceno); Paramo Mucubaji, camino a Mucunuque, 3600 m, 25 Oct 1980, B. Briceno & Adamo 244 (Herbarium Briceno, US); 4200^1300 m, 4 Nov 1992, Meier 3025 (VEN); Distr. Campo Elias, Sierra La Culata, Pico El Campanario, 4325m, 20 Oct 1972, Teran 7732 (MERF); 3600-4100 m, 26 Oct 1972, Teran 7909 (MERF); 18. Festuca flmbriataNees, Fl. Bras. Enum. PI. 2(1): 472.1829. (Figs. 22,23,82A-E). TYPE: Brasilia meridionales, Montevideo, F. Sellows.n. (holotype: B!; isotypes: K!, LE-TRIN-2806.01!, LETRTN-2806.02!,US-557541exB!,US-1441522 exB!, US-1126679 fragm. ex W!, W!). Festuca ampliflora Doll, Fl. Bras. 2(3): 116, pi. 34. 1878. TYPE: Brazil. Minas Gerais, Caldas, Regnell III 1409 (lectotype: S! designated by Alexeev, Bot. Zhurn. (Moscow & Leningrad) 69: 348. 1984; isolectotypes: P!, US-2875375 fragm. ex M!). Rhizomatous perennials with extravaginal innovations. Culms (60-)90-150(-250) cm tall, erect, glabrous; nodes 3 or 4 in distal half; cataphylls sometimes present, about 4-6 cm long. Leaf sheaths membranous, brown, striate, glabrous; auricles absent; ligules 0.1•0.5(•1) mm long, coriaceous, apex truncate, ciliate; blades 20-50 x 0.4-1.5 cm, flat, green, abaxially scabrous with prickles on abaxial epidermis. Panicles 20^t0 x 10-15 cm, ovate; branches scabrous. Spikelets 11-15(-20) mm long, oblong or ovate, florets (5-)6-7(-9); rachillas 1-1.3 mm long, glabrous; glumes (3.5-)4-8(-9) mm long, lanceolate, scabrous, apex acute; lower glumes (3.5-)4-6(-7) mm long, 1-nerved; upper glumes 5.5-6(-7.5) mm long, 3-nerved; lemmas 6-8 (-9) mm long, membranous to coriaceous, lanceolate, 5-nerved, scabrous, papillose, apex entire, awnless; callus glabrous; paleas as long as the lemma, scabrous; lodicules ca. 1mm long, lanceolate, with scattered hairs at apex; anthers 2.5-3.5(-4) mm long; ovary apex pilose. Caryopses lanceolate; hilum nearly as long as the grain. Leaf blade anatomy.•Cross-sections with numerous vascular bundles with small ribs above; sclerenchyma under both abaxial and adaxial epidermis, discontinuous and extending to the vascular bundles; bulliform cells present; epidermis with scattered prickles. Observations.•Festuca fimbriata is characterised by having pilose lodicules. This unusual feature probably represents a plesiomorphy since elsewhere in the genus it is only known to occur in the Australian F. subg. Austrofestuca. Distribution and habitat.•Festuca fimbriata is restricted to the area between northern Uruguay, northeastern Argentina, and southeastern Brazil, in forest openings, wet depressions, and swamps between (70-)700-1650(-2400)m. Additional specimens examined. ARGENTINA. Corrientes: Dept. Santo Tome, Ea. Timbo, costa del Rio Uruguay, 26 km SE de Colina Garabi, 16 Sep 1980, O. Ahumada & A. Schinini 4085 (CTES); Arroyo Chimiray y Ruta 40, 8 Oct 1976, Quarin 3409 (CTES); Dept. Santo Tome, Garruchos, 22 Oct 1954, Burkart 19654 (BAA); Estancia "Garruchos", swamps and wet medows, T.M. Pedersen 819 (C, MO); Ruta 37, 5 km E de Gdor. Virasoro, 14 Nov 1974, A. Schinini & Carnevali 10523 (US). Misiones: Dept. Capital, Ruta provincial No. 1,12kmS de Posadas, 16 Nov 1974, A. Schinini & Carnevali 10687 (CTES). BRAZIL. Minas Gerais: Jan 1880, Regnells.n. (W); Caldas, 28 Jan 1846, Widgren s.n. (K, US); Monte Verde, Sep 1997, Longhi & Witten 5011 (ICN). Parana: Festuca in South American Paramos 53 Figure 22. Festucafimbriata. A. Stylized growth form. B. Habit. C. Inflorescence. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Lodicule. J. Leaf blade cross-section. A-J, T.M. Pedersen 11475 (MO). 54 Festuca in South American Paramos -80 -71) -50 -f.l) -40 -30 • •* » t Li* - - . '. II) II) \A PANAMA VENEZUELAN £ /-'Amu•' MAS \ ( \s*+LJ V ECJJADORf-Tgk. -10 \ PERU BRAZIL -21) -21) -30 y I f -80 -70 / / *y • {{\ A -M\ 0 500 1000 Kilomet N \y -60 -50 -40 -31 Figure 23. Distribution of Festuca fimbriata (•), and F. andicola (A). Piraquara, Pinhais,, 25 Nov 1968, G. Hatschbach 20379 (C, K); Mun. Balsa Nova, Bicudo, 6 Dec 1962, G. Hatschbach 9551 (US); 26 Feb 1909, Dusen 8972 (K, US); Mun. Curitiba, Atuba, margin of swamp, 23 Jan 1965, Clayton 4301 (K); Castro, Rio Cunhaporanga, varzea, 26 Nov 1968, G Hatschbach 20381 (K); Mun. Jaguariaiva, Rio Jaguariaiva, campo, 13 Nov 1974, G. Hatschbach 35575 (K). Rio de Janeiro: Mun. Itatiaia, Prateleiras, 2300 m, Mar 1937, Brade & Kuhlman 15631 (B, COL, PRC, RB); Serra de Itatiaia, 1900 m, Mar 1894, Ule 243 (R); Rio de Janeiro, Alto de Itatiaia, 2200-2400 m, 17 Jan 1925, A. Chase 8302 (US). Rio Grande do Sol: Aparados da Rocinha, Bom Jesus, 18 Jan 1950, Rambo 45452 (US); Vacaria, 9 Jan 1946, J.R. Swollen 8189 (MO, US); Mun. Rio Pardo, Fasenda Soledad (Jurgens), 70 m, 1905, Jurgensen G85 (US); Bom Jesus, entre Ausentes e Poste Fiscal da Serra da Rocinha, 4 Dec 1971, Vails et al. 1893 (US); Facenda Carauna-Bom Jesus, Dutra 543 (R); Mun. Julio de Castilhos, between Abacatu & Venado, 13 Nov 1976, T.M. Pedersen 11475 (C, CTES, K, MO, P); Mun. Vacaria, Jul 1954, Barreto 577 (BAA); S. Luiz, 24 Nov 1952, Rambo 53471 (B, US); Faz Englert Sao Francisco de Paula, 1 Jan 1952, Rambo 54704 (B, US); Serra de Rocinha, Bom Jesus, 1200 m, 18 Jan 1950, Rambo 45466 (B); 3 Feb 1953, Rambo 53816 (B, US); Cambara do Sul, Itaimbezinho, 15 Dec 1973, Hitchcock 28 (ICN); Taimbezinho, Nov 1977, Boechat 41059 (ICN); Cambara do Sul, Itaimbezinho, 7 Jan 1977, Sampaio & Arzivenco 341, 362 & 404 (ICN); Bom Jesus-Faz. Arechavaleta, Cavauna, Jan 1903, Dutra 538 (ICN); Bom Jesus-estr. Para Vacaria, 15 km apos ponte de saida de B. Jesus, 30 Nov 1975, Longhi & Sampaio 368 (ICN). Santa Catarina: Mun. Mafia, Campo Novo, 7 km from Mafra, 9 Dec 1971, Smith & Klein 15761 (ICN, K, R, US); Campo Novo, Mafra, 750 m, 12 Dec 1962, Klein 3872 (K, US); Mun. Porto Uniao, 45 km S, Festuca in South American Paramos 900-1100 m, 5 Feb 1967, Smith & Klein 10855 (US); Campo de Areao, Santa Cecilia, 1100 m, 19 Dec 1962, Reitz & Klein 14172 (US); Mun. Cacador, 8 km N of Cacador, 950-1100 m, 7 Feb 1957, Smith & Klein 10958 (US); Mun. Sao Joaquim, source of Rio Capivaras, Serra do Oratorio, 10 km E of Bom Jardim da Serra (Cambajuva), 1200 m, 16 Jan 1957, Smith & Reitz 10127 (US); Mun. Bom Retire, Fazenda Campo dos padres, 1650 m, 25 Jan 1957, Smith & Reitz 10428 (US); Mun. Marombas, Curitibanos, Banhado, 900 m, 9 Jan 1962, Reitz & Klein 11812 (US); Base of Morro Juco Prudente, 1 Jan 1946, J.R. Swollen 8089 (MO, US); Mun. Bom Jardim de Serra, 10 km S of Bom Jardim at Rio Capivaras, 15 Dec 1971, Smith & Klein 15805 (K, MO, R, US); Mun. Curitibanos, 28 Nov 1971, Smith & Klein 15474 (K, R, US); Mun. Lajes, 8 km S of Painel, 19 Dec 1971, Smith & Klein 15916 (K, R, US); 27 km S of Lajes, 20 Dec 1971, Smith & Klein 15936 (K, R, US); Mun. Chapeco, campo, bog and pinheiral, Fezenda Campo Sao Vicente, 24 km W of Campo Ere, 900-1000 m, 26 Dec 1956, Smith et al. 9389 (R, US); Mun. Ere, 6 km W of Campo Ere, 26°22'S, 53°06'W, 900-1000 m, 6 Dec 1964, Smith & Klein 13674 (K, P, R, US); Campo de Chatorio, Ule 616 (W); Walde am Capirone, Serra Geval, Mar 1891, Ule 1959 (P); Sao Biento, 22 Jun 1885, Schwaike s.n. (P); Mun. Rio Pardo, 70 m, Nov 1908, Jurgens 175 (W); Campos Dos Padres, 22 Jan 1957, Rambo 60082 (B); Curitibanos, Banhadoi, 900 m, 10 Jan 1962, Reitz & Klein 11948 (B, US). Sao Paulo: Campos do Jordao, in paludosis, 1600 m, Feb 1946, Leite 3490 (US); Mun. Sao Roque, 50 km W from, Sao Paulo, 850 m, 12 Feb 1987, Tsugaru & Otsuka B-2265 (MO). PARAGUAY. Caazapa: Tavai, bosque cercano al hospital, 28 Oct 1988, Degen 861 (FCQ, MO); galery forest 1 km N of hospital, 26°10'S, 55°27'W, 28 Oct 1988, Zardinii 7701 (FCQ, MO). Itapua: Cordill. San Rafael, Estero Yukeri, Bernardi 18618 (MO). URUGUAY. Caticeiras Rivera, 11 Dec 1907, Berro 5994 (BAA). Tacuarembo. Campos de Tacuarembo, terrenos arenosos, Arechavalet s.n. (W); Montevideo, 1826, D'Orbygny 39 (P); Cerro Largo, Cerro de Las Cuentas, 29 Dec 1938, Rosengurtt B-2764 (US). 19. Festuca andicola Kunth, Nov. Gen. Sp. (quarto ed.) 1: 153.1816. (Figs. 23, 24, 83C & D). Festuca racemosa Willd. ex Spreng., Syst. Veg. 1: 352. 1825, nom. inval. TYPE: Ecuador. Carchi, Paramo de Puntas, 3220 m, Humboldt 55 & Bonpland s.n. (holotype: P!; isotypes: B!, P!, US-2875373 fragm. ex P!). Perennials with well developed rhizomes, innovations extra vaginal. Culms 50-70 cm tall, erect, solitary, glabrous; nodes 2 or (3) in basal half. Leaf sheaths membranous, brown, striate; auricles absent; ligules 0.5-0.9 mm long, membranous to coriaceous, apex truncate, ciliate; blades 10-17 x 0.1-0.4 cm, flat or conduplicate, green, abaxially glabrous. Panicles 12-17 x 0.5-1 cm, contracted, erect, elongate; branches mostly glabrous. Spikelets 7-9 mm long, florets 3 or 4(-5); rachilla glabrous or with scattered hairs; glumes 1.4-2.5 mm long, membranous to coriaceous, oblong-lanceolate, purplish, apex obtuse ciliate; lower glumes 1.4-1.8 mm long, 1-nerved; upper glumes 2-2.5 mm long, 3-nerved, apex short-ciliate; lemmas 5-5.5 mm long, 5-nerved, membranous to coriaceous, lanceolate, purplish, upper third scabrous, apex acute or short-awned, the awn or mucro 0.5-0.7 mm long; callus glabrous; paleas as long as the lemma, membranous, glabrous, scabrous on keels, apex hairy; lodicules lanceolate, acuminate; anthers 0.8-1.1 mm long; ovary apex glabrous. Caryopses not seen. Leaf blade anatomy.•Cross-sections with 7•9(•11) vascular bundles and 5-7(-9) ribs above; sclerenchyma under abaxial epidermis discontinuous, some fascicles extending to the vascular bundles forming girders; adaxial epidermis with scattered hairs, the hairs 0.05-0.09 mm long. Observations.•Saint-Yves (1927, 180) mentions Festuca dissitiflora Steud., a nom. nud., as belonging to F. andicola. Examination of the voucher of F. dissitiflora housed at US (W. Lechler 1829, US-2875397 fragm. ex GOET!) confirms that this species has nothing to do with F andicola and that it is a synonym of F rigescens (J. Presl) Kunth. Historically, the name F andicola had been applied to a heterogeneous group, and this group was divided into two species by Stancik (2003) where a new type and name, F soukupii, were designated. Festuca andicola and F soukupii are morphologically similar to the Venezuelan species, F elviae, and the Peruvian F. tenuiculmis Tovar. Festuca tenuiculmis differs from the rest of the group by having ramified, opened panicles. Festuca andicola differs from the others by having solitary culms with rhizomes, whereas F soukupii is caespitose forming small bunches. Distribution and habitat.• Festuca andicola ranges from southern Ecuador to northern Colombia. Tovar (1972) mentions F. andicola as being 56 Festuca in South American Paramos 2 mm Figure 24. Festuca andicola. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf blade cross-section. A-G, Stancik 3551 (PRC). Festuca in South American Paramos present in Peru but no Peruvian specimens were found during preparation of this revision. It occurs in swamps and margins of streams in the grass paramo zone and Andean mountain forests between 2900-3 800(-4000) m. This species is associated, in addition to many others, with communities of the order Marchantio-Epilobietalia (Cleef 1981). Additional specimens examined. COLOMBIA. Antioquia: Mun. Urrao, Paramo Frontino, La Cueva, 3450 m, 11 Sep 1986, Rolddn et al. 396 (COL, HUA, MO); Llano Grande, 3400 m, 10 Sep 1986, Rolddn et al. 307 (HUA). Boyaca: Mun. Aquitania, Paramo Los Curies, 3500 m, 8 Feb 1999, D. Stancik 2190 (COL, FMB, PRC); Mun. Mongui, Paramo de Laguna La Colorada, 3600 m, 21 Jan 1999, D. Stancik 2024 (COL, PRC); Mun. Santa Rosa, Paramo Alto Lamadero, 3200 m, 30 Feb 1988, D. Stancik 1455 (COL, PRC); Paramo de Pisba, carratera Socha-La Punta, 3570 m, 11 Jun 1972, Cleef 4376 (COL, U, US); Mun. Arcabuco, way from Laguna Iguaque to Laguna Ojo de Agua, 3650 m, 15 Oct 1998, D. Stancik 948 (COL, FMB, PRC); Mun. Toca, Paramo Cortadero, 5°30'N, 73°15'W, 3350 m, 14 Nov 1998, D. Stancik 1365 (COL, FMB, PRC). Caldas: Mun. Manizales, Parque Nacional Los Nevados, 4020 m, 18 Sep 1999, D. Stancik 3412 (COL, PRC); way from Rio Nereidas to Casa del Cisne, km 5-7, 4000 m, 18 Sep 1999, D. Stancik 3399 (COL, PRC); Los Nevados, Villamaria, 3 Feb 1985, Safin & Herndn 1060 (FMB). Cauca: Macizo Colombiano, Valle de las Papas, alrededores de Valencia, 2910 m, 11 Sep 1959, Idrobo et al. 3678 (COL). Cundinamarca: Mun. San Juan de Sumapaz, Parque Nacional Sumapaz, Alto de Toquilla, 3800 m, 15 Nov 1999, D. Stancik 3554 (AAU, COL, PRC); 15 Nov 1999, D. Stancik 3551 (COL, PRC); road from San Juan to Usme, km 5-7, 3650 m, 15 Nov 1999, D. Stancik 3547 (COL, PRC); Parque Nacional Chingaza, mina Palacio, 3700 m, 11 Oct 1992, Figueredo 119 (COL, HPUJ); 3700 m, 5 Aug 1992, Figueredo 126 (HPUJ); Muncipio Fomeque, Valle de los Frailejones, 3150 m, 12 Jun 1989, Bernal & Jimenez 1127 (HPUJ); Monserrate, Aug 1859, Lindig 14 (P); Alto San Juan, 4000 m, R. Fosberg 20749 (US). Meta: Paramo de Sumapaz, Hoya El Nevado, Laguna La Guitarra, 3405m, 24 Jan 1973, Cleef 8285 (COL). Nariflo: Mun. Tuquerres, Volcan Azufral, road from Vereda San Roque to Laguna Verde, km 3,2650 m, 9 Mar 1999, D. Stancik2775 (COL, PRC, PSO); El Encano, vereda Catapamba, 2900 m, 23 Mar 1999, D. Stancik 2983 (AAU, 57 COL, PRC, PSO, W); Cumbal, 3300 m, 16 Apr 1986, Benavides 6497 (COL, PSO). Norte de Santander: Mun. Chilos, vereda El Hatico on road Pamplona-Bucaramanga, 3500-3700 m, 1 Dec 2000, D. Stancik (COL, PRC). Risaralda: Laguna del Otun, quebrada Juntas, 3750 m, 29 Aug 1985, Sanchez & Hernandez 950 (FMB). Santander: Paramo de Berlin, cerca del Picacho, 3290 m, 29 Sep 1966, Robinson & Beltran 3142 (US). ECUADOR. Carchi: road Las Juntas-El Angel, km 14, 3400 m, 11 Mar 1992, S Laegaard 101731 (AAU, QCA, QCNE). Chimborazo: Mun. Riobamba, Volcan Chimborazo, sector Cruce de Los Arenales, 01°28'14"S, 78°54'06"W, 4300 m, 20 Sep 2000, D. Stancik 3712 (PRC, QCA); D. Stancik 3705 (AAU, PRC, QCA). Cotopaxi: Volcan Cotopaxi, road to Limpiapungo, 00°37'S, 78°27'W, 3850 m, S. Laegaard 5866 (AAU, QCA). Imbabura: Cordillera Oriental, Paramo de Angochagua, 29003600 m, Acosta-Solis 18836 (US); Mun. Urcuqui, road to Cerro Yanaurcu, 00°28' 13"N, 78° 18 '45"W, 4100 m, 15 Oct 2000, D. Stancik 4092 (PRC, QCA). Pichincha: Lloa, 3000 m, E. Asplund 7524 (AAU, F, S); Paramo de Mojanda-LagunaNegra, 00°08'S, 78°16'W, 3800 m, 14 May 1985, S Laegaard 54327 (AAU, QCA, QCNE); Road Pifo-Papallacta, 3 km W of Paso de la Virgen, 00°18'S, 78°14'W, 37003900 m, 7 Aug 1985, S. Laegaard 54892 (AAU, QCA); Cerro Ungui, 2860 m, Hartman 7a (US); Mun. Amiguana, Volcan Pasachoa, 00°29'51"S, 78°29'25"W, 3700 m, 14 Sep2000,D. Stancik3685 (PRC, QCA); Mun. Amaguana, Volcan Pasachoa, 00°30 '23"S, 78°29 '28"W, 3350 m, 14 Sep 2000, D. Stancik 3683 (AAU, PRC, QCA). Tungurahua: Cotalo, 2900 m, Acosta-Solis 9879 (US); Slope of Mt. Tungurahua above Banos, 2800 m, E. Asplund 8427 (NY, S). 20. Festuca rubra L., Sp. PI. 1: 74. 1753. (Figs. 21,25,83A & B). TYPE: In paludosis prati regii Upsalia, Anonymous (lectotype: GB!, designated by Jarvis et al., Watsonia 16:302. 1987.) Loosely tufted and rhizomatous perennials with extravaginal (rarely intravaginal) innovations. Culms 20-50 cm tall, erect, glabrous; 1 node in basal half. Leaf sheaths membranous to coriaceous, brown, striate, glabrous or pilose, margins united at base; auricles absent, ligules 0.3-0.5 mm long, membranous to coriaceous; blades 7-15 cm long, 0.7-1 mm wide, conduplicate to involute, green, sometimes glaucous, abaxially glabrous, apex Festuca in South American Paramos 58 2 cm 2 mm Figure 25. Festuca rubra. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf blade cross-section. A-G, Stancik 3457 (PRC). Festuca in South American Paramos obtuse. Panicles ca. 10 cm long, 3-5 cm wide, erect or spreading; branches scabrous. Spikelets 10-12 mm long, florets 5 or 6; rachilla glabrous or with scattered hairs; glumes 4-5.5 mm long, membranous to coriaceous, lanceolate, purple, apex acute; lower glumes ca. 4 mm long, 1-nerved; upper glumes 5-5.5 mm long, 3-nerved; lemmas 6.5-7 mm long, 5-nerved, membranous to coriaceous, lanceolate, green to purplish, apex scabrous, awned, the awn 2.5-3.5 mm long; callus glabrous; paleas almost as long as the lemma, scabrous on keels, apex hairy; lodicules oblong-lanceolate; anthers 3.5-3.7 mm long; ovary apex glabrous. Caryopses not observed. Leaf blade anatomy.•Cross-sections with 7 vascular bundles and 5 ribs above; sclerenchyma discontinuous under both abaxial and adaxial epidermis, not forming girders; adaxial epidermis with scattered hairs, the hairs 0.02 mm long. Distribution and habitat.•This species is introduced from Europe and rarely cultivated. Additional specimens examined. COLOMBIA. Caldas: Mun. Neira. Vereda Chupaderos, Sector Campamentos, pasture, 3650 m, D. Stancik 3457 (COL, PRC). ECUADOR. Chimborazo: Paramo de Urbina, 25 km N of Riobamba, ca. 2 km W of Panamerican Hwy, 3500 m, 28 Jan 2000, S. Laegaard et al. 20994 (AAU, LOJA, QCA, PRC). Pichincha: Picalqui al S de Tabacundo, 2750 m, Acosta-Solis 16358 (F, US). 21. Festuca soukupii Stancik, Folia Geobot. Phytotax. 39(1): 103, f. 2, 1-5. 2004. (Figs. 26, 27, 83E & F). TYPE: Ecuador. Imbabura, Mun. Cayambe, Volcan Cayambe, 00°31.6'N, 78°55.6'W, swamps below the refuge with Loricaria sp., Festuca glumosa, Distichia muscoides Nees & Meyen, etc., 4450 m, 20 Dec 2000, D. Stancik 4162 (holotype: PRC!; isotypes: AAU!, QCA!). Small tussocked perennials with intra- or extravaginal innovations. Culms 10-50 cm tall, erect, glabrous; nodes l(-2) in basal half. Leaf sheaths membranous, brown, striate, densely hairy and appearing shaggy, margins free; auricles absent; ligules 0.3-0.5 mm long, membranous to coriaceous, apex obtuse, short-ciliate; blades 5-25 cm long, 0.5-0.8 mm wide, conduplicate to involute, green, abaxially glabrous, apex obtuse. Panicles 2.5•13(•17) cm long, 0.3-0.5 cm wide, erect, contracted, elongate; branches hairy. Spike- 59 lets 6-8(-9) mm long, florets 3 or 4(-5); rachilla glabrous or with short scattered hairs; glumes 1.2-2.5(-2.7) mm long, membranous to coriaceous, ovate to lanceolate, purplish, apex obtuse, ciliate; lower glumes 1.2-2 mm long, 1-nerved; upper glumes 2-2.5(-2.7) mm long, 3-nerved, apex shortciliate; lemmas 4.5-5.5 mm long, membranous to coriaceous, lanceolate, 5-nerved, purplish, papillose, apex awned, sometimes scabrous, the awn (0.5-)0.7-1.7 mm long; callus glabrous; paleas as long as the lemma, glabrous, scabrous on keels, apex hairy; lodicules oblong-lanceolate; anthers 0.8-1.1 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 3/5 of total length. Leafblade anatomy.•Cross-sections with 5-7 vascular bundles and 3-5 ribs above; sclerenchyma under abaxial epidermis discontinuous, under adaxial epidermis absent [larger plants with 7•9(•11) vascular bundles and adaxial schlerenchyma, sometimes forming 2-A girders]; adaxial epidermis with scattered hairs, the hairs 0.03-0.05 mm long. Observations.•Festuca soukupii is morphologically similar to F. andicola. However, F andicola has well developed rhizomes (versus tussock forming) and glabrous sheaths (versus densely hairy). Distribution and habitat.•Festuca soukupii ranges from the Cordillera Central of northern Colombia, through Ecuador to northern Peru. It is known from humid grass paramo and super-paramo vegetations between 3700-4500 m. In addition, this species is found in large tufts along trails and roads up to the forest zone between 2600-3200 m. Additional specimens examined. COLOMBIA. Caldas: Mun. Manizales, Parque Nacional Los Nevados, Hda. La Esperanza, 3500-3700 m, 18 Sep 1999, D. Stancik 3416 (COL, PRC); Mun. Neira, vereda Chupaderos, sector "Campamentos", 3650 m, 26 Sep 1999, D. Stancik 3455 (COL, PRC); Paramo de Quindio, 3700-4200 m, 20 Aug 1922, Pennell 9951 (US). Cauca: Mun. Popayan, Parque Nacional Purace, Pilimbala, 02°22.1'N, 76°24.06'W, 3250 m, 6 Jul 2000, D. Stancik 3610 (COL, PRC); Macizo Colombiano, Valle de las Papas, alrededores de Valencia, 2910 m, 11 Sep 1958, Idrobo et al. 3682 (COL, US). Nariflo: Mun. Cumbal, Volcan Nevado de Cumbal, NE slopes, 3700 m, 9 Mar 1999, D. Stancik2744 (AAU, COL, PSO, PRC); Mun. El Encano, between vereda Catapamba and road Pasto-El Encano, 2800 m, 23 Mar 1999, D. Stancik 2981 (COL, PSO, PRC); Plain N of Guachucal, 3000 m, 1 Mar 1986, Wood 60 Festuca in South American Paramos 1 mm Figure 26. Festuca soukupii. A, B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A, C-G, Stancik 3887 (PRC); B, Stancik 3079 (PRC); H, Stancik 4162 (PRC). Festuca in South American Paramos -80 -75 -70 61 -65 Figure 27. Distribution of Festuca soukupii (•), F. chitagana (A), and F. hatico (•). 5331 (COL, K). Risaralda: Mun. Pereira, Parque Nacional Los Nevados, below Nevado del Cisne, way from Casa del Cisne to Laguna de Otun, km 5, 4100 m, 19 Sep 1999, D. Stancik 3380 (PRC, W); below Nevado Santa label, crossroads to Villa Maria, Hda. Potosi and Casa del Cisne, 40004100 m, 20 Sep 1999,/). Stancik3373 (COL, PRC, US). Tolima: N side of Nevado del Ruiz, 3800 m, 19 Feb 1984, Wood 4247 (COL, FMB, K); Mun. Ibague, Nevado del Tolima, La Cueva, 04° 39'N, 76°19.5'W, 3900 m, 8 Jun 2000, D. Stancik 3596 (COL, PRC). ECUADOR. Azuay: Parque Nacional Cajas, slopes near Laguna Luspa, 3700 m, 21 Apr 1990, P.M. Peterson 8858, C.R. Annable & M.E. Poston (QCA, QCNE, US); along Rio Miguir, 3480 m, 21 Apr 1990, P.M. Peterson 8870, C.R. Annable & M.E. Poston (QCA, QCNE, US); around lake Llaviaucu, 02°56'S, 79°10'W, 3400 m, Oxford Expedition 30 (K); El Pan, mountain zone, 2800 m, Harlington 1197 (S); around Lagoon TaglacochaTres Cruces, 02°47'S, 79°13'W, 3900-4000 m, 1 Sep 2000, D. Stancik 3858 (PRC, QCA); road Cuenca-Saraguro, km 42, 03°10'S, 79°02'W, 3450 m, 19 Feb 1985, S Laegaard 53580 (AAU, QCA); road Cuenca-Limon, towards antennas, 03°10'S, 79°02'W, 3350 m, 21 Nov 1999, S Laegaard et al. 20824 (LOJA, AAU); Mun. Nabon, antennas on the road Loja-Cuenca, 03°12'S, 79°02'W, 3300-3450 m, 30 Aug 2000, D. Stancik 3802, 3808 (QCA, PRC). Bolivar: pass on road Guaranda-Riobamba, 01°35'S, 78°50'W, 40504150 m, 10 Jul 1990, S. Laegaard 71737 (AAU, QCNE); highway to Guaranda km 66.5 and to Fecundo km 2.2, 4300 m, 03 May 1990, P.M. Peterson 8996 & C.R. Annable (MO, QCA, QCNE, US); Simiatung, Hacienda Talahua, 3400 m, Penland & Summers 583 (F). Canar: road CanarChunchi, 02°24'S, 78°59'W, 3200 m, 05 Mar 1985, 62 Festuca in South American Paramos S. Laegaard 53837 (AAU, QCA, QCNE); El Tambo, road El Tambo-Ingapirca, 3400-3500 m, D. Stancik 3800 (PRC, QCA). Carchi: km 21.2 NE of El Angel and 27.8 km SW of Tulcan, above Rio Bolo, 3320 m, 14 May 1990, P.M. Peterson 9139, E.J. Judziewicz & R.M. King (MO, QCA, US); km 11.4 NE of El Angel on road to Tulcan, 3240 m, P.M. Peterson 9118, E.J. Judziewicz & R.M. King (US); km 14 along road Las Juntas-El Angel, 3400 m, 11 Mar 1997, S. Laegaard 101718 (AAU, QCA, QCNE). Cotopaxi: road ZumbahuaPujili, km 39, 00°53'S, 78°47'W, 3750-3800 m, S. Laegaard 102087 (AAU); Paramo de Laguna Salayampe, E of Latacunga, 00°56'S, 78°25'W, 4000^1100 m, S. Laegaard 54148 (AAU); Volcan Cotopaxi, N slope, 00°39'S, 78°26'W, 4200 m, Sklendf & Kosteckova 1905 (AAU); Parque Nacional Cotopaxi, Laguna Limpiopungo, 00°40'S, 78°30'W, 3800 m, 3 Apr 1982, Fegan & Falconi 9 (QCA); Falda NNW, 4180 m, 03 Jul 1986, Ehrenburg 60 (QCA); Valley NW of Limpiapungo, 00°38'S, 78°28'W, 3900^000 m, 25 Feb 1992, S. Laegaard 101432 (AAU, QCA); N slope on the left side of the road to refuge, 00°39'S, 78°26'W, 4200 m, 19 Jul 1995, Sklendf & Kosteckova 85-8, 86-9, 87-6 (NY); Quebrada ca. 3 km de carratera, 00°40'S, 78°30'W, 3400 m, 08 May 1982, Bravo 56, 58 (QCA); railway station Cotopaxi, 3400 m, E. Asplund 6478 (S, US); Cotopaxi, 2500 m, E. Asplund 6400 (S); road Salcedo-Napo, km 2739,4000-4100 m, Sparre 15708 (S); Mun. Chaupi, NE slope of Volcan Illiniza, 37'59"S, 78°42'42 "W, 4000^1050 m, 12 Oct 2000, Z>. Stancik 4023, 4028, 4031 (PRC, QCA); Mun. Lasso, volcan Cotopaxi, 00°39'6"S, 78; 78°30'55"W, 3530 m, 28 Sep 2000, D. Stancik 3108, 3128, 3879, 3887 (AAU, PRC, QCA). Chimborazo: Volcan Chimborazo, E side, 01°28'S, 78°46'W, 4260 m, Sklendf & Sklendfovd 2262 (AAU); 4300 m, Sklendf & Sklendfovd 2168 (AAU); km 42.7 SW of Ambato on Highway to Guaranda, 4020 m, P.M. Peterson 8973 & C.R. Annable (MO, QCA, QCNE, US); km 10 E of Lago Colta on road to Pallatango, 3725m, P.M. Peterson 9209, E.J. Judziewicz, R.M. King & P.M. Jorgensen (K, MO, QCA, QCNE, US); between Urbina and Mt. Chimborazo, 3600^500 m, A.S. Hitchcock 21981 (US); 3750 m, E. Asplund 7892 (K, S); 3900 m, E. Asplund 8447 (P, S); Paramo de Urbina km 25 N of Riobamba, ca. 2 km W of Panamerican Hwy, 01°29'S, 78°42'W, 3500 m, S. Laegaard et al. 20993 (AAU); S slope of Mt. Chimborazo, 3800 m, Fagerlind & Wibon 934-bis (S); km 9 NE of San Juan de Velasco on road to Lago Colta, 3600 m, 21 May 1990, P.M. Peterson 9235, E.J. Judziewicz, R.M. King & P.M. Jorgensen (QCA, QCNE, US); Daldal Valley km 10 E of Licto, 01°48'S, 78°32'W, 3700 m, Ramsay etal. 104A (K); 00°15'S, 78°28'W, 3450 m, 19 Aug 1987, Ramsay & Smith 245, 247 (K, QCA, QCNE); along Rio Alao, 01°52'S, 78°30'W, 3200-3400 m, S. Laegaard 55301 (AAU); near pass between Volcan Chimborazo and Carihuairazo, 01°27'S, 78°48'W, 4400 m, S. Laegaard & Sanchez 20027 (AAU); Carihuasco above Mashahuasca, 01°29'S, 78°49'W, 4500 m, S. Laegaard & Sklendf 20363 (AAU); road Guamote-Macas km 15, 02WS, 78°40'W, 3750 m, S. Laegaard & Sklendf 20335 (AAU, LOJA); Mun. Riobamba, volcan Chimborazo, sector Cruce delosArenales,01°27'51"S,78°53'58"W, 4150 m, 20 Sep 2000, D. Stancik3713 (PRC, QCA). Imbabura: Hacienda Mojanda on road to Otavalo, 2900-3000 m, Sparre 13516 (AAU, S); Laguna Grande, 00°08N, 78°16'W, 3725-3750 m, S. Laegaard 52374A (AAU, QCA); Laguna Cuicocha, 3100 m, E. Asplund 20217 (S); Mun. Urcuqui, road to Cerro Yanaurcu, 00°26'28"N, 78°15'24"W, 4100 m, 15 Oct 2000, D. Stancik 4090B, 4093 (PRC, QCA); Mun. Cayambe, volcan Cayambe, 00°00'26.5"N, 78°01 '21"W, 4300 m, 20 Oct 2000, D. Stancik 4158,4162 (PRC, QCA); Mun. Urcuqui, road to Cerro Yanaurcu, 00°37'46"N, 78°41 '45"W, 4100 m, 15 Oct 2000, D. Stancik 4094 (PRC, QCA). Loja: km 10 along road to Fierra Urcu, 03°41'S, 78°18'W, 3400 m, S. Laegaard et al. 19066 (AAU, LOJA, QCA); Cerro de Arcos W of road Manu-Zaruma, 03°34'S, 79°28'W, 35003600 m, S. Laegaard & Aguirre 20609 (AAU); Paramo de Carboncillo, km 11 S of Ona, 2800 m, S. Laegaard 19739 (AAU, QCA); Mun. Saraguro, road to Fierra Urcu, 03°41'S, 79°16'W, 30003100 m, 24 Aug 2000, D. Stancik 3787,3 780 (PRC, QCA, US). Morona-Santigo: Hda. HuarguallaHda. San Eduardo, way to Parque Nacional Sangay, 02°00.25'S, 78; 27'W, 3700 m, 20 Jul 1999, D. Stancik 3320 (PRC, QCA); Parque Nacional Sangay, Plaza Culebrillas, 01°58'S, 78°25'W, 3500-3600 m, 22 Jul 1999, D. Stancik 3358, 3359 (PRC, QCA). Napo: Valle Vicioso E of Volcan Cotopaxi, 3600 m, Holm-Nielsen & Balslev 23755 (AAU); El Tambo SE of Volcan Cotopaxi, 00°42'S, 78°18'W, 3650 m, S Laegaard 55535 (AAU). Pichincha: road Pifo-Pintag, 2.5 hours horseride above Inga Moserat, 00°19'S, 78°17'W, 3950 m, S Laegaard 102268,102237(AAU, QCA); Paramo Festuca in South American Paramos de Guamani, 3960 m, 23 Nov 1991, Leon 1230 (QCA); 4200-4250 m, S. Laegaard 101384 (AAU); 4300 m, S. Laegaard & S.A. Renvoize 70519 (AAU); 4100 m, S. Laegaard 103107 (AAU, QCA); road Quito-Santo Domingo near San Juan, 00°17'S, 78°37'W, 3450-3500 m, S. Laegaard 52605,52607 (AAU, QCA); Andinum Quitensium Pichincha, Jameson 296 (P, W); from Pichincha, 4000ft, Jun 1859, Jameson 14 (W); Quito-Panecillo, 2900 m, E. Asplund 6020 (S, US); Mun. Otavalo, shrubby margin and pajonal on the road from Otavalo to Laguna Mojanda, 00°09'54"N, 78°17'20"W, 3450m, 19 Oct 2000,D. Stancik4111 (PRC, QCA); Nevado Cayambe, W side of the volcano, 00°01'N, 78°01'W, 4200 m, Sklendr & Kosteckova 66-10 (US); road to Refuge, 4300 m, S. Laegaard & S.A. Renvoize 70519 (AAU); Paramo de Trujillo between peaks of Illinizas and Cotopaxi, 3450 m, Barclay & Juajibioy 7961 (MO, US); Tunel de agua Quito-Papallacta, 00°22'S, 78°08'W, 16 Mar 1994, Aguirre & Merino 4219 (AAU, LOJA); E slope of Illiniza Sur, 4300 m, Sklendr & Kosteckova 16-1 (US); 4200 m, Sklendr & Kosteckova 15-5 (NY); Volcan Illiniza, N slope, 4000-4100 m, Sparre 15635 (S); road Lloa-Guagua Pichincha, km 10, 00°13'S, 78°35'W, 4170 m, Laegaard et al. 102737 (AAU); Rucu Pichincha, 4400 m, E. Asplund 17313 (S); NE slope, 4300 m, Sklendr & Kosteckova 1-24 (NY); 4600 m, E. Asplund 8606 (S); slope above Lloa, 3200 m, E. Asplund 7558 (S); WNW slope of Antisana, 4600 m, Halloy B-54 (AAU); between La Libertad and San Juan, 3200 m, E. Asplund 16259 (S); Canal on W side of Volcan Atacazo, 00°20'S, 78°38'W, 3750-3800 m, S. Laegaard 55673 (AAU, QCA); slopes of Ruminahui, 3900 m, Sparre 15871 (S); Mun. Pifo, paramo de Guamani, 00°19'S, 78°15'W, 3700 m, 19 Jun 1999, D. Stancik 3004 (AAU, PRC, QCA); Mun. Amaguana, Volcan Pasachoa, 00°30'23"S, 78°29'28"W, 3350 m, 14 Sep 2000, D. Stancik 3686 (PRC, QCA). Tungurahua: km 10 S ofMocha, 3600 m, Hading et al. 10533 (GB); Cotalo, 2900 m, Acosta-Solis 9884 (US); AcostaSolis 9877 (US); Mt. Carihuayrazo, 4400 m, E. Asplund8470 (S); Mun. Pillaro, Las Llanganatis, around Aucacocha lagoon, 01°8'55"S, 78°20'00.4"W, 3800 m, 28 Sep 2000, D. Stancik 3903 (PRC, QCA). Zamora-Chinchipe: Road Loja-Zamora, ca. 2-6 km E of pass, 03°59'S, 79°09'W, 2600 m, S. Laegaard 18736 (AAU, LOJA, QCA, QCNE). PERU. Cajamarca: 52 km N of Cajamarca on Hwy 3N towards Bambamarca, 63 small lagoon with Ranunculus and open grasslands with Deyeuxia, 3780 m, 16 Mar 2000, P.M. Peterson 14907 & N. Refulio Rodriguez (MO, US, USM); Prov. Chota, Miracosta, entre Miracosta y Pampa del Lirio, 3380 m, 11 Dec 2000, Vega et al. 10320 (F, MO); Pampa Larga, al N de la explotacion minera Yanacocha, 3900 m, 14 May 1994, Vega 7143 (F); Jalca de Kumullca, ruta a Celendin, 3650 m, 7 Feb 1975, Vega et al. 1638 (F); Cerro Sexcemayo, al W de Cajamarca, jalca graminosa, 3500 m, 4 Feb 1991, Vega 5419 (F); Prov. Celendin, Cajamarca-Celendin road, especially near large rock outcrops, 3000-3450 m, 28 May 1984, Smith etal. 7314, 7321a, 7332 (MO); Celendin, Jalca de Kumulca, mountain pass on the road CajamarcaCelendin, 07°02'31"S, 78°15'33"W, humid jalca vegetation with tussock grasses, 3700 m, 26 Aug 2004, Sklendr & Cruz 8706 (PRC). 22. Festuca azucarica E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 69:1546.1984. (Figs. 28, 29, 84A-D). TYPE: Colombia. Valle de Cauca, Cordillera Central, cerro Pan de Azucar, 3700 m, 26 Feb 1969, J. Cuatrecasas, Espinal & Ramos 27562 (holotype: US-278!; isotypes: COL!,U-5508!). Dense tussock forming perennials with intravaginal innovations. Culms 70-80 cm tall, erect, glabrous; nodes 1 or 2 in distal half; cataphylls 0.5-1.5 cm long, short, coriaceous. Leaf sheaths coriaceous, brown, glabrous; auricles absent; ligules 0.3•1(•1.5) mm long, coriaceous, apex truncate, short-ciliate; blades 30-50 cm long, 0.9-1.4 mm wide, conduplicate, rigid, olive-green, abaxially glabrous, apex mucronate. Panicles 10-25 x 1•3(•6) cm, lanceolate to ovate, mostly contracted; branches glabrous or finely scabrous. Spikelets 9-10 mm long, lanceolate, florets 4-6; rachilla densely hairy; glumes (2.5-)3-5(-6) mm long, coriaceous, narrowly lanceolate, dark purple, sparsely scabrous; lower glumes (2.5-)3-3.5(-4) mm long, 1-nerved; upper glumes 4-5(-6)mm long, 3-nerved; lemmas (5.5-)6-7.5 mm long, 5-nerved, membranous to coriaceous, lanceolate, dark purple, scabrous and densely hairy, apex entire, awnless or short-awned, the awn 0.3-0.5 mm long; callus sparsely hairy; paleas as long as the lemma, papillose, upper third hairy; anthers 2.5-3.5 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 1/2-3/5 as long as the grain. Leaf blade anatomy.•Cross-sections with 9-15 vascular bundles and 7-14 ribs; sclerenchyma 64 Festuca in South American Paramos Figure 28. Festuca azucarica. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 3377 (PRC). Festuca in South American Paramos 65 BRAZIL 0 200 400 600 Kilometers -80 -75 -70 -65 Figure 29. Distribution of Festuca colombiana (•), F. dasyantha (A), F. azucarica (•), and Flaegaardii (•). under abaxial epidermis continuous or discontinuous, united with all the vascular bundles, adaxial sclerenchyma present, extending to all vascular bundles; bulliform cells absent; epidermis densely hairy adaxially, the hairs 0.4-1.3 mm long. Observations.•Festuca azucarica is morphologically similar to F. pilar-franceii, a species endemic to the Colombian Cordillera Oriental. However, F. azucarica differs by having short catophylls 0.5-1.5 cm long (versus 2-5 cm in F pilarfranceii) and longer lemmas (5.5-)6-7.5 mm long (versus 4.5-6 mm). Festuca dasyantha, F colombiana, F. laegaardii, and F. toca are all species in F. sect. Cataphyllophorae that are characterized by the presence of cataphylls. Distribution and habitat.•Festuca azucarica is endemic to the Colombian Cordillera Central (Caldas, Quindio, Risaralda, Tolima, and Valle de Cauca). It is one of the dominant species of the matorral zone and grass paramo and occurs between 3500-4000 m. This species is found in many different plant communities that contain Calamagrostis recta (Kunth) Trin. ex Steud., Festuca aff sublimis & Diplostephium rupestre (Kunth) Wedd. (Cuatrecasasl934), Espeletietum hartwegianaeCalamagrostiosum effusum (Cuatrecasas 1934), Festuco dolichophyllae-Calamagrostietum effusae (Salamanca-V. 1991) and Espeletia hartwegiana subsp. centroandina Cuatrec. and Calamagrostis recta (Cleefetal. 1983). Additional specimens examined. COLOMBIA. Caldas: Mun. Manizales, Parque Nacional Los Nevados, way from Casa del Cisne to Rio Nereidas, km 5, 3800-4000 m, 18 Sep 1999, D. Stancik 3404, 3405 (COL, PRC); Nevado El Ruiz Libano, 4000 m, 21 Jul 1958, Barclay 6431 (COL, US); 4500 m, Aug 1984, Yepes-Agredo 760 (COL); 3400-3500 m, 17 Dec 1917, Pennell 3003 66 Festuca in South American Paramos (US); Nevado de Santa Isabel, Quebrada de Leon, 3800 m, 24 Nov 1946, J. Cuatrecasas 23137 (U, US, VALLE); Cabeceras del Rio Otiin, Laguna Taburetes, 3580 m, 24 Nov 1946, J. Cuatrecasas 23178 (US, VALLE); Nevado del Ruiz, 8 Oct 1983, Wood 4033 (AAU, K); Parque Nacional Los Nevados, 5 May 1940,/. Cuatrecasas 9292 (COL). Quindio: Mun. Salento, vereda Cocora, below Nevado del Quindio, 22 May 1989, J. Luteyn et al. 13051 (COL); Pijao, Paramo de Chili, 3600 m, 17 Sep 1998, Correa etal. 129 (COL); Mun. Salento, Paramo de Romerales, 3680 m, 30 Oct 1994, Velez etal. 4514 (HUQ). Risaralda: Mun. Pereira, Parque Nacional Los Nevados, below Nevado Santa Isabel, crossroads to Villa Maria, Casa del Cisne and Hda. Potosi, 4000-4100 m, 20 Sep 1999, D. Stancik 3375, 3376, 3377 (COL, PRC); Mun. Pereira, Parque Nacional Los Nevados, 3510 m, 27 Jul 195$,Jaramillo-Mejia & Cleef5 738 (COL); Exped. Botan. Novae-Granatae, sin. loc, Mutis 5545, 5555 (MA, US). Tolima: Mun. Santa Isabel, Quebrada Africa, 3900 m, 19 Feb 1980, Jaramillo-Mejia et al. 6181 (COL); 3800 m, 10 Feb 1980, Diaz-Piedrahita & Rangel 2037 (COL); Los Valles, tinea La Cascada, cabeceras del Rio Anaime, 4000 m, 10 Feb 1980, Echeverry 1973 (COL, TOLI); Mun. Ibague, Nevado del Tolima, around La Cueva, 04°39'N, 75°19.5'W, grass paramo, 3900 m, 7-8 Jun 2000, D. Stancik 3599 (COL, PRC). 23. Festuca chitagana Stancik, Darwiniana 41(1^): 130, f. 12g-k. 2003. (Figs. 27, 30). TYPE: Colombia. Santander, Mun. Cerrito, paramo del Almorzadero, km 15 on the road from Cerrito to Chitaga, 3700 m, 25 Feb 1999, D. Stancik & Medina 2577 (holotype: PRC!; isotypes:COL!,FMB!). Rhizomatous perennials forming small tussocks with intra- and extravaginal innovations. Culms 60-80 cm tall, erect, glabrous; nodes 2-A in distal half; cataphylls short, coriaceous, graybrown; Leaf sheaths membranous to coriaceous, brownish-gray, glabrous, striate; auricles absent; ligules 0.5-1 mm long, membranous to coriaceous, apex truncate, ciliate; blades 15-20 cm long, 0.7-0.9 mm wide, involute, abaxially glabrous, olive-green, apex obtuse. Panicles 10-12 cm x 9-12 cm, triangular, flexuous, nutant, branches glabrous. Spikelets 6-9 mm long, lanceolate, florets 2 or 3; rachilla sparsely short-hairy; glumes 4-6.5 mm long, membranous, glabrous, apex acute; lower glumes 4-5 mm long, narrowly lanceolate, 1-nerved; upper glumes 5.5-6.5 mm long, oblonglanceolate, 3-nerved; lemmas 6-7 mm long, 5-nerved, lanceolate, membranous to coriaceous, hairy, apex acute or short-awned, the awn 0.5-1 mm long; callus glabrous; paleas 4/5 as long as the lemma, lanceolate, membranous, densely hairy; lodicules 0.8-1 mm long, lanceolate; anthers 1-1.6 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 4/5 as long as the grain. Leafblade anatomy.•Cross-sections with 7-9 vascular bundles and 5-7 ribs above; sclerenchyma under abaxial epidermis discontinuous; adaxial sclerenchyma and girders absent; adaxial epidermis sparsely hairy, the hairs ca. 0.07 mm long. Observations.•Festuca chitagana is morphologically similar to the Mesoamerican species F talamancensis Davidse that is known only from Costa Rica. However, F talamancensis has shorter lower glumes (3.5-3.8 mm versus 4-5 mm in F. chitagana), shorter upper glumes (5-5.3 mm versus 5.5-6.6), and an oblong caryopsis (versus lanceolate) with a hilum only 3/5 as long as the grain (versus 4/5). Distribution and habitat.• This species is endemic to Colombia and is known only from the northern Cordillera Occidental (Paramo de Frontino) and Oriental (Paramo del Almorzadero, Sierra Nevada del Cocuy). It occurs in the grass paramo zone with shrubs mostly on stony substrates between 3400^000 m. Additional specimens examined. COLOMBIA. Antioquia: Mun. Urrao, Paramo de Frontino, Churumblum-La Mosca, 3400 m, 14 Nov 1984, X. Londono et al. 550 (COL, MEDEL). Boyaca: Mun. El Cocuy-Guican, Parque Nacional El Cocuy, Las Cabanas Kanwara, way to Pico Ritacuba along river Playita, 4070 m, D. Stancik & Carvajal 1866 (PRC). 24. Festuca colombiana E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 69: 1546, f. 2, 7-8. 1984. (Figs. 29, 31, 84E-F, 85A-B). TYPE: Colombia. Cundinamarca, Paramo de Sumapaz, Chisaca, Laguna Negra, orilla sur, pedregal humedo con Sphagnum sp., 3800 m, 11 Dec 1971, A.M. Cleef181 (holotype: US2785658!; isotypes: COL!, P!, U!, VEN!). Short rhizomatous perennials forming small tussocks with intra- and extravaginal innovations. Culms 40-80(-120) cm tall, erect, finely scabrous; nodes 1 or 2(-3) in distal half; cataphylls small Festuca in South American Paramos 67 2 mm Figure 30. Festuca chitagana. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik & Carvajal 1866 (PRC). 68 Festuca in South American Paramos Figure 31. Festuca colombiana. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 1889 (AAU). Festuca in South American Paramos (about 1-2 cm long), coriaceous. Leaf sheaths membranous to coriaceous, glabrous, stramineous to grayish, margins free; auricles absent; ligules 0.1-0.3(-0.5) mm long, coriaceous, apex truncate, short-ciliate; blades (15-)20-30(-40) cm long, 0.4•0.6(-0.8) mm wide, conduplicate to involute, rigid, abaxially glabrous, green, apex acute. Panicles 10-20(-25) x 1-2 cm, contracted; branches densely short-villose. Spikelets 9-12 mm long, oblong-lanceolate, florets 4•6(-7); rachilla densely pilose; glumes 3•5.5(•6.5) mm long, lanceolate, coriaceous, purplish, upper 1/4 scabrous, apex acute; lower glumes 3•4(•4.5) mm long, 1-nerved; upper glumes 4.5-5.5(-6.5) mm long, 3-nerved; lemmas (5.5-)6-7(-7.5) mm long, 5-nerved, lanceolate, membranous to coriaceous, purplish, papillose, upper 1/3 sparsely scabrous, apex entire, mucronate or awned, the awn up to 2 mm long; paleas as long as the lemma, papillose, upper 1/3 hairy; lodicules ca. 1.1 mm long, lanceolate, acuminate; anthers 2-3.5 mm long; ovary apex glabrous. Caryopses oblonglanceolate; hilum 2/3 as long as the grain. Leaf blade anatomy.•Cross-sections with 79(•13) vascular bundles and 5-11 ribs; sclerenchyma under abaxial epidermis discontinuous, rarely continuous, extending to all the vascular bundles, adaxial sclerenchyma extending to every second vascular bundle; bulliform cells absent; adaxial epidermis densely hairy, the hairs 0.02-0.09 mm long. Observations.•This species has a relatively large area of distribution and plants from different localities are morphologically variable. It is morphologically similar to F. azucarica and F pilar-franceii, as mentioned previously. Distribution and habitat.•Festuca colombiana is endemic to Colombia where it is known only from the Cordillera Oriental (Boyaca, Cundinamarca, Meta, Santander). It can be found growing in grass paramos between 3200-4000 m. This species is known from the following vegetation communities: Espeletio argentae-Arcytophylletum nitidi (Rangel & Ariza 2000), Diplostephio phylicoidis-Espeletietum grandi-florae (Rangel & Ariza 2000), Calamagrostietum planifoli-effusae (Vargas & Zuluaga 1985), Espeletio argentaeCalamagrostiosum effusum (Cuatrecasas 1934), and Bartsio santolinifoliae-Calamagrostietum effusae (Rangel & Ariza 2000). Additional specimens examined. COLOMBIA. Boyaca: Mun. Arcabuco, Santuario Iguaque, Laguna Iguaque, 3450-3500 m, 15 Oct 1998, D. Stancik 928 (COL, FMB, PRC); Mun. Saboya, 69 Paramo Saboya, 3000 m, 24 Oct 1998, D. Stancik 1065,1069 (CO, FMB, PRC); Mun. Aquitania, Paramo Sarna, 3400 m, 20 Jan 1999, D. Stancik 1969 (COL, FMB, PRC); D. Stancik 1978 (COL, PRC); 3100m, 14 Jan 1999,D. Stancik 1923{COL, FMB, PRC); D. Stancik 2003 (COL, PRC); Mun. Aquitania, Paramo Los Curies, 3500 m, 8 Feb 1999, D. Stancik2168, 2192 (COL, FMB, PRC); 3500 m, 8 Feb 1999, D. Stancik 2174, 2218 (COL, FMB, PRC); Mun. Samaca, Paramo Rabonal, 200 m, 1 Nov 1998, D. Stancik 1323, 1324, 1325 (COL, FMB, PRC); D. Stancik 1297 (COL, PRC); vereda Ruchcal, 1 Nov 1998,/). Stancik 1287,1288,1290, 1302,1319,1321 (COL, FMB, PRC); Mun. Aquitania, road Aquitania-Sisvaca, km 6,2900 m, 10 Feb 1999, D. Stancik2256(COL, FMB, PRC); 3100 m, 13 Jan 1999, D. Stancik 1924 (COL, FMB, PRC, W); 3350 m, 14 Jan 1999, D. Stancik 1921 (COL, FMB, PRC); Mun. El Cocuy, Parque Nacional Nevado del Cocuy, Las Cabanas Kanwara, 3950 m, 30 Dec 1998, D. Stancik 1875 (COL, FMB, PRC); paramo on the road to Pico Ritcuba, km 2-3 from Cabanas, 4000 m, 30 Dec 1999, D. Stancik & Carvajal 1811, 1848,1891 (COL, FMB, PRC); D. Stancik & Carvajal 1874 (COL, PRC); El Cocuy-Guican, Parque Nacional El Cocuy, sector Lagunillas, road from Casa Herrera to lagoons, km 3-4, 3950 m, 30 Dec 1998, D. Stancik 1851, 1854 (COL, FMB, PRC); Hac. La Esperanza-Valle de los Frailejones, 3750 m, 30 Dec 1998,£>. Stancik & Carvajal 1783, 1787 (COL, FMB, PRC); Hac. La Esperanza, rocky slope in the front of house, fields and rest of the Polylepis forest, 3650 m, 30 Dec 1999, D. Stancik 1842 (COL, FMB, PRC); Mun. Mongui, Laguna La Colorada, 3550 m, 21 Jan 1999, D. Stancik 2025 (COL, FMB, PRC); Mun. Toca, Paramo Cortadero, 05°30'N, 73°15'W, 3300 m, 14 Nov 1998, D. Stancik 1352, 1355, 1357, 1391, 1402 (COL, FMB, PRC); Mun. Paipa, cuchilla El Paramo, 3200 m, 3 Dec 1998, D. Stancik 1512, 1516,1530,1531, 7546 (COL, FMB, PRC); NW of Duitama, Paramo de La Rusia, 3400 m, 2 Jul 1984, Wood 4479 (COL, K). Cundinamarca: El Chico, Bogota, Dec 1946, Black 46-605 (NY); Mt. Guadalupe above Bogota, bushy slope, 2800-3300 m, 9 Dec 1917, Pennell 1923 (NY); Paramo de Guasca, entre Guasca y Gacheta, 3320 m, 1 Jun 1959, Barclay & Juajibioy 6517 (MO); Paramo entre Cogua y San Cayetano, cercania de la Laguna Seca, 3700 m, 12 Nov 1972, Cleef6258 (COL, US); Chapinero near Bogota, 3000-3100 m, 23 Sep 1917, Pennell 2018 (MO, US); Paramo de Monserrate, 3300 m, 70 Festuca in South American Paramos 12 Sep 1987, Sanchez 283 (COL); Parque Nacional Sumapaz, Laguna Chisaca, 3700-3800 m, 10 Feb 1961, Pinto-Escobar & Hernandez 523 (COL); 3706 m, Pedraza etal. 491 (COL, PRC); Cuchilla LaRabona, 4020 m, 11 Feb 1972, Cleef1587 (COL, U); Mun. Usme, Laguna Negra, 3750 m, 20 Feb 1986, Rangel & Aguirre 3720 (COL); Paramo de Chisaca, 3910 m, 11 Nov 1958, Barclay &Juajibioy 6109 (COL, MO, US); 3650-3700 m, 29 Oct 1959, J. Cuatrecasas & Jaramillo-Mejia 25742 (COL); Laguna Negra, 3800 m, 11 Dec 1971, Cleef 181 (COL, P, U, US); 3500 m, 16 Jul 1998, D. Stancik 303 (COL, PRC); Mun. Guatavita, vereda Carbonel Alto, 3200 m, 30 Oct 1999, D. Stancik 3500 (COL, PRC); Guadalupe, 3000 m, Jul 1913, Apollinaire & Arthur 121 (US); Mun. Bogota, Alto de la Viga, 3550 m, 1 Nov 1999, D. Stancik 3538, 3540, 3542 (COL, PRC); Mun. Tausa, cuchilla Los Cuervos-Laguna Verde, 3650 m, D. Stancik 3496 (COL, PRC); Mun. Choachi, Paramo de Cruz Verde, laguna Verjon, 3450 m, 19 Oct 1985, Castellanos et al. 3 (BOG); 3720 m, Aug 1935, Garcia-Barriga 1016,1019 (AAU, COL); Choachi, 2900 m, Lindig 1013 (K, P); 2900-3200 m, Aug 1859, Lindig 1053 (US); 3200 m, 18 Dec 1915, Apollinaire & Arthur 9 (US); Paramo de Palacio, 2 km al lado de Mina, 3750 m, 29 Nov 1972, Cleef & Urribe 6694 (COL, U, US); Paramo de Guasca, El Santuario, 3200 m, 26 Jan 1972, J. Cuatrecasas 3550 (MA); Mun. San Juan de Sumapaz, road to Usme, km 5-7, 2950 m, 1 Nov 1998, D. Stancik 1289 (COL, PRC); 15 Nov 1999, D. Stancik 3544, 3545, 3546, 3560 (COL, PRC); Mun. Santa Rosa de Viterbo, Paramo Alto Lamadero, Laguna Sagrado Corazon, 3200 m, 30 Nov 1998, D. Stancik 1436 (COL, FMB); Mun. Siachoque, paramo Siachoque, 5°28'51"N, 73°12'44"W, 23 Jan 1999, D. Stancik 2073 (COL, PRC); Las Tronaderas, 3770 m, 24 Jan 1999, D. Stancik 2063 (COL, PRC); vereda Carnichoque Arriba; 3400 m, 24 Jan 1999, D. Stancik 2053 (COL, PRC); 23 Jan 1999, D. Stancik 2052 (PRC, COL); 1998, D. Stancik 2408 (COL, PRC); Vereda San Antonio, 3600 m, 16 Oct 1999, D. Stancik 3491, 3492, 3493, 3494 (COL, PRC); cuchilla El Muchacho, 3600 m, 16 Oct 1999, D. Stancik 3480 (COL, FMB, PRC); Mun. Zipaquira, paramo Guerrero, 3500 m, 16 Oct 1999, D. Stancik 3481 (COL, FMB, PRC); Paramo de Chisaca, way to Chisaca, near Rio Tunjuelito, 3750 m, 20 Sep 1966, T.R. Soderstrom 1229 (COL, K, TULV); Paramo El Tablazo, frequent in recently burned paramo, 3200 m, 2 Apr 1983, Wood 3612 (K). Meta: Macizo de Sumapaz, Hoya de la Quebrada Clarincito, Los Frailes, 3720 m, 2 Jul 1981, DiazPiedrahita 2367 (COL). Santander: Mun. Cerrito, Paramo del Almorzadero, vereda Mortyno-La Cascada, 3400 m, 25 Feb 1999, D. Stancik 2527, 2550,2562, 2563, 2564,2565 (COL, FMB, PRC); 25 Feb 1999, D. Stancik 2560, 2561 (COL, PRC); Mun. Conception, vereda Juradito, Paramo de Gallina, 3200 m, 24 Feb 1999, D. Stancik 2487, 2507, 2508, 2511, 2512 (COL, FMB, PRC); Vicinity of Vetas, rocky hillside, 3100-3250 m, 20 Jan 1927, Killip 17329 (US); Exped. Botan. Novae-Granatae, sin. loc, Mutis 5570 (MA). 25. Festuca dasyantha Kunth, Nov. Gen. Sp. (quarto ed.) 1: 154-155. 1816. (Figs. 29, 32, 85C & D). TYPE: Ecuador. Cotopaxi, in Montis Cotopaxi, 4090 m, Humboldt & Bonpland s.n. (holotype: P!; isotypes: B!, BAA-1187 fragm. ex B!, B-W-2071!, US-2875395 fragm!, W!). Tufted perennials with short rhizomes and intravaginal innovations. Culms 40-100 cm tall, erect, glabrous; nodes 2 or 3 (-4) in basal half; cataphylls 1-4 cm long, coriaceous, dark brown, striate. Leaf sheaths membranous to coriaceous, brown, striate, glabrous or scabrous; ligules 0.3-0.5 mm long, membranous to coriaceous, margins ciliate, apex truncate; blades 15-35 cm long, 0.71.4(-2.2)mm wide, mostly conduplicate, green, abaxially glabrous. Panicles 15-30 x 1-3 cm, narrow, erect, branches glabrous or sparsely scabrous. Spikelets 10-11 mm long, florets 4 or 5; rachilla densely pilose; glumes 3-5 mm long, lanceolate, membranous to coriaceous, green, distal 1/3 often densely hirsute, apex acute; lower glumes 3-3.5 mm long, 1 -nerved; upper glumes 4.5-5 mm long, prominently 3-nerved; lemmas 5.5-6 mm long, lanceolate, 5-nerved, membranous, green, densely hairy, apex entire, acute or short-awned, the awn, up to 1 mm long; paleas as long as the lemma, glabrous, upper 1/3 hirsute; lodicules lanceolate, acuminate; anthers 2.3-3.3 mm long; ovary apex glabrous. Caryopses not seen. Leaf blade anatomy.•Cross-sections with 9-17 vascular bundles; sclerenchyma discontinuous under both abaxial and adaxial epidermis, extending to every other or nearly all vascular bundles forming girders; bulliform cells not observed; adaxial epidermis densely hairy, the hairs 0.7-1.3 mm long. Observations.•Festuca dasyantha differs from all other species in F sect. Cataphyllophorae, Festuca in South American Paramos 71 Figure 32. Festuca dasyantha. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-H, Laegaard 101260 (AAU); I, Mille 664 (QPLS). 72 Festuca in South American Paramos such as F.procera, F. colombiana, and F laegaardii, by having glumes and lemmas bearing long hairs. Distribution and habitat.•Festuca dasyantha is endemic to central Ecuador (Carchi, Imbabura, Pichincha). It occurs in grass paramos between 2800-3000 m. Additional specimens examined. ECUADOR. Carchi: Road el Angel-Mira, km 3, 00°37'N, 77°58'W, 2950 m, S. Laegaard 101260 (AAU, QCA). Imbabura: Entre Yuquin y Cebadal, Sigsipamva, 2800 m, Acosta-Solis 18729 (US). Pichincha: Pifo, pascuis andinis et subandinis, 3000 m, Mille 664 (QPLS); 2700 m, Mille 659 (QPLS); between Pifo and Los Coralles, 2800 m, E. Asplund 6978 (K, NY, S, US); Quito-Panecillo, Sodiro s.n. (MO, NY, QPLS, US). Note.• Specimen Mutis 5554 (US ex MA) comes from Ecuadorian collection of F.J. Caldas and dating to the period of Mutis's stay in Colombia. 26. Festuca hatico Stancik, Darwiniana41(l^t): 119, f. 121-p. 2003. (Figs. 27, 33, 86A & B). TYPE: Colombia. Norte de Santander, Mun. Chilos, Vereda Hatico on road Bucaramanga-Pamplona, 3500-3700 m, 1 Dec 2000, D. Stancik 4290 (holotype: PRC; isotype: COL). Tussocked perennials with rhizomes and intraand extravaginal innovations. Culms 60-70 cm tall, erect, glabrous; nodes 2 or 3 in distal half; cataphylls coriaceous, gray-brown, striate. Leaf sheaths coriaceous, brownish-stramineous, glabrous, striate; auricles absent; ligules 0.3-0.5 mm long, membranous to coriaceous, bilobed; blades 30-40 cm long, 0.7-0.9 mm wide, conduplicate to involute, abaxially glabrous, green, apex obtuse. Panicles 10-20 x 0.5-1 cm, contracted, narrow; branches finely scabrous. Spikelets ca. 9 mm long, lanceolate, florets 3 or 4; rachilla short, papillose; glumes 3.2-5 mm long, coriaceous, glabrous, apex acute; lower glumes 3-3.6 mm long, narrowly lanceolate, 1-nerved; upper glumes 4.5-5 mm long, lanceolate, 3-nerved; lemmas 6-6.5 mm long, 5-nerved, membranous to coriaceous, oblong-lanceolate, papillose, apex scabrous and two-dentate, awned, the awn 0.3-0.7 mm long; callus glabrous; paleas 4/5 as long as the lemma, lanceolate, membranous, papillose, upper 1/2 hairy; lodicules ca. 1.7 mm long, triangular; anthers ca. 3.5 mm long; ovary apex glabrous. Caryopses not seen. Leaf blade anatomy.•Cross-sections usually with 9 vascular bundles and 7 ribs above; sclerenchyma under abaxial and adaxial epidermis discontinuous, extending to some vascular bundles forming girders; adaxial epidermis hairy, the hairs, 0.4-0.9 mm long, often numerous. Observations.•Festuca hatico is morphologically similar to Festuca colombiana. However, the latter species has wider panicles (1-2 cm versus 0.5-1 cm), spikelets with 4-7 florets (versus 3 or 4), pilose rachillas (versus papillose), and paleas as long as the lemmas (versus 4/5 as long). Distribution and habitat.•Festuca hatico is endemic to the northeastern Andes and is known from Colombian Cordillera Oriental (Norte de Santander, Santander) and Venezuela (Lara). It can be found growing in shrubby patches of the paramo zone between 3100-3900 m. Additional specimens examined. COLOMBIA. Norte de Santander: Mun. Chilos, Vereda Hatico on road Bucaramanga-Pamplona, 3500-3700 m, 1 Dec 2000, D. Stancik 4290 (COL, PRC). Santander: East slope of Paramo de las Coloradas, above La Baja, 3900 m, 27 Jan 1927, Killip 18479 (US). VENEZUELA. Lara: Mun. Humocaro Alto, Parque Nacional Dinira, 9°35'39"N, 70°07'12"W, 3170 m, D. Stancik 4289 (CAR, COL, PRC). 27. Festuca laegaardii Stancik, Folia Geobot. Phytotax. 39(1): 107, f. 4, 6-10. 2004. (Figs. 29, 34,86C-F). TYPE: Ecuador. Tungurahua/Napo, Mun. Pillaro, Las Llanganatis, 01°09'37.5"S, 78°14'50.8"W, Valle de los Frailejones, margins of the swamp dominated by Carex sp. and dryer shrub formation, 3500 m, 28 Sep 2000, D. Stancik 3983 (holotype: PRC!; isotypes: AAU!, QCA!). Rhizomatous and loosely caespitose perennials with extravaginal innovations. Culms (15•)30•100 cm tall, erect, glabrous; nodes 2 or 3(-5) in distal half; cataphylls present, short, membranous, brown, striate. Leaf sheaths membranous, purplishbrown, striate, hairy; ligules l-1.5(-2) mm long, membranous to coriaceous, apex truncate, ciliate; blades 20-25 x 0.5-1.1 cm, conduplicate, abaxially glabrous, rarely hairy. Panicles (5•)10•15(•18) x 1-2 (-11) cm, compressed, rarely flexuous, branched; branches densely hairy. Spikelets (7.5-) 8-10(-12) mm long, florets (3-)4 or 5(-6); rachilla sparsely pilose; glumes 2.8-5.5(-6) mm long, coriaceous, narrowly lanceolate, purple, upper 1/3 densely hairy, apex acute; lower glumes 2.8-3.7 (-4) mm long, 1-nerved; upper glumes 4.5-5.5 Festuca in South American Paramos 73 1 mm mm «' 1 cm 2 mm Figure 33. Festuca hatico. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 4290 (PRC). 74 Festuca in South American Paramos 1 mm Figure 34. Festuca laegaardii. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 3983 (PRC). Festuca in South American Paramos (-6) mm long, 3-nerved; lemmas 5.5-6.5 mm long, 5-nerved, lanceolate, membranous to coriaceous, green-purple, apex acute, entire, upper 1/3 hairy; paleas almost as long as the lemma, glabrous or inconspicuously scabrous, apex hairy; lodicules obovate; anthers 2.4•2.8(-3) mm long; ovary apex glabrous. Caryopses not observed. Leaf blade anatomy.•Cross-sections with 7-11 vascular bundles and 3-7 ribs above; sclerenchyma under both abaxial and adaxial epidermis discontinuous, extending to some vascular bundles forming girders; bulliform cells not observed; adaxial epidermis usually with scattered hairs, the hairs 0.45-0.7 mm long. Observations.•Festuca laegaardii is morphologically similar to Festuca dasyantha. However, the latter species has densely hairy adaxial epidermis(versus scattered hairs in F. laegaardii), lemmas densely hairy throughout (versus only short hairy on the upper third), shorter ligules (0.30.5 mm long), and glabrous panicle branches. Distribution and habitat.•Festuca laegaardii is endemic to Ecuador and is known only from Azuay, Canar, Carchi, and Napo. This species occurs in humid and swampy patches in grass paramos between 2900-3400 m. Additional specimens examined. ECUADOR. Azuay: Road Gualaceo-San Juan Bosco, ca. 3 km from pass, 03°00'S, 78°39'W, 3250-3350 m, S. Laegaard 53965 (AAU, QCA); Road Cuenca- Saraguro, km 68, 03°20'S, 78°10'W, 3350 m, S. Laegaard 53597 (AAU, QCA). Canar: Road BiblianCanar, 3400 m, Harling et al. 8635 (GB). Carchi: Alor-Hda. San Rafael, 2900-3000 m, Acosta-Solis 21031 (US). Morona-Santiago: Parque Nacional Sangay, Plaza Culebrillas, 01°58'S, 78°25'W, forest grassy patches, 3500-3700 m, D. Stancik 3365 (PRC, QCA). Napo: Laguna San Marcos NE of Volcan Cayambe, 00°07'N, 77°58'W, 3370m,B. Ollgaard et al. 34039 (AAU, PRC); Papallacta-NE de la laguna principal, 3400 m, Jaramillo et al. 11855 (QCA); Jaramillo et al. 11856 (MO, QCA). 28. Festuca pilar-franceii Stancik, Darwiniana 41(1-4): 121, f. 14a-e. 2003. (Figs. 35, 37, 87A & B). TYPE: Colombia, Cundinamarca, Mun. Bogota D.C., sector Santa Rosa-Laguna Chisaca, 3500-3650 m, 7Aug 1998,/). Stancik 3585 (holotype: PRC!; isotype: COL!). Densely tussocked perennials with intravaginal innovations. Culms 70-100(-120) cm tall, erect, 75 glabrous; nodes 1 or 2 in distal half; cataphylls 2-5 cm long, membranous to coriaceous. Leaf sheaths membranous to coriaceous, brown-stramineous, glabrous; auricles absent; ligules 0.1-0.3 mm long, coriaceous, apex truncate; blades 30-60 cm long, 0.7-1.1 mm in diameter, conduplicate to involute, abaxially glabrous, green, apex obtuse. Panicles (12-)15-20(-25) x 1-7 cm, oblonglanceolate, branched, flexuous, erect; branches glabrous or finely scabrous. Spikelets (8•)9•11 mm long, lanceolate, florets 4•6(-7); rachilla densely pilose; glumes 2-5 mm long, narrow lanceolate, coriaceous, glabrous, purple or purplish-green, apex acute; lower glumes 2-3(-3.5) mm long, 1-nerved; upper glumes 3-5 mm long, 3-nerved; lemmas 4.5-6 mm, lanceolate, 5-nerved, membranous to coriaceous, purplish-green, scabrous or hairy, apex mucronate or short-awned, the awn < 1 mm long; paleas as long as the lemma, papillose, upper 1/3 hairy; lodicules lanceolate, acuminate; anthers 2.5-3.5 mm long; ovary apex sparsely hairy. Caryopses not seen. Leafblade anatomy.•Cross-sections with 1115 vascular bundles and 9-13 ribs; sclerenchyma under abaxial epidermis continuous, extending to all the vascular bundles forming girders; bulliform cells absent; adaxial epidermis densely hairy, the hairs 0.8-1.2 mm long. Observations.•Festuca pilar-franceii is morphologically similar to F. azucarica, a species from the Cordillera Central. In the Cordillera Oriental, F cleefiana is another species that is morphologically similar to F. pilar-franceii. However, F. cleefiana lacks cataphylls, has longer ligules [(0.5-)l-2(-2.5) mm], and has longer lemmas (6-8 mm). Distribution and habitat.•This species is endemic to the Colombian Cordillera Oriental (Macizo de Bogota, Paramo de Sumapaz). It occurs in the swampy patches and margins of lagoons in grass paramos between 3600-4000 m. Festuca pilar-franceii is associated with two communities: Baccharis revoluta & Cortaderia cf. nitida (Sturm & Rangel 1985) and Chusquea tesellata, Espeletia grandiflora, and Calamagrostis effusa (Sturm & Rangel 1985). Additional specimens examined. COLOMBIA. Cundinamarca: Cabrera, Paramo de Cruz Verde, 3720 m, VIII 1935, Garcia-Barriga 1018 (COL); Sumapaz, a 2 km de la division de carretera a San Juan, 3960 m, 3 Oct 1978, Rangel 1650 (COL); km 51-60 via San Juan, 10 Feb 1986, Torres-Romero & Lozano 2982 (COL); Laguna 76 Festuca in South American Paramos Figure 35. Festucapilar-franceii. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 315 (PRC). Festuca in South American Paramos Chisaca, 3625m, 25 Aug 1972, Cleef 5259 (COL, U, US); Paramo Chisaca, 3750-3960 m, Oct 1966, T.R. Soderstrom 1307 (US, TULV); near headwater of Rio San Juan, 18 km E of Cabrera, 4°5'N, 74°12'W, 4000 m, 11 Aug 1943, R. Fosberg20745 (US); Macizo de Bogota, Laguna Negra, 3720 m, 11 Sep 1961, J. Cuatrecasas & Jaramillo-Mejia 25865 (COL, US); 3600-3700 m, III 1973, Cleef 3605(COL,P,U); 16 Jul 1998,/). Stancik210, 217, 218, 224, 225, 226, 231, 241, 248, 250, 256, 257, 258, 264, 265, 267, 268, 270, 277, 278, 302, 311, 312, 313, 314, 315, 316, 317, 318, 319, 320, 321 (COL, PRC); 9 Aug 1999, D. Stancik 3559, 3562, 3563, 3564, 3565, 3569, 3568, 3579, 3580, 3583, 3584, 3586, 3587, 3589, 3590 (COL, PRC); Mun. Bogota, Laguna Chisaca, 9 Aug 1999, D. Stancik 3566, 3567, 3570, 3572, 3573, 3574, 3575, 3576, 3577, 3578, 3581, 3582, 3585, 3588 (COL, PRC); 6 Jan 1997, D. Stancik 491 (COL, PRC); 15 Nov 1999, D. Stancik 3559 (COL, PRC, US); 7 Aug 1998, D. Stancik 228 (COL, PRC); D. Stancik 3494 (COL, PRC); D. Stancik3561 (COL, PRC); Parque Nacional, Sumapaz, vereda Santa Rosa, ladera de la quebrada Bijuacales, localidad 20, 3500 m, 07 Aug 1998, Pedraza et al. 228, 242, 278 (COL, PRC); Paramo de Sumapaz, between St. Rita and San Juan, 3600 m, 15 Jul 1984, Wood 4539 (K, US). Meta: Paramo Sumapaz, Hoya de Quebrada Sitiales, 0.5 km al SW de Laguna La Primavera, 3550 m, 26 Jan 1972, Cleef1048 (COL, U); Cerro Nevado de Sumapaz, 3615m, 29 Jan 1972, Cleef 1272 (COL); lado SW, 3590 m, 12 Jan 1973, Cleef 7704 (COL). 29. Festuca procera Kunth, Nov. Gen. Sp. (quarto ed.) 1: 154. 1816. (Figs. 36, 37, 87C-F). Diplachne procera (Kunth) Spreng., Syst. Veg. 1: 351. 1825. Festuca orgyalis Willd. ex Spreng., Syst. Veg. 1: 351. 1825, nom. inval. Festuca scabra Willd. ex Steud., Nomencl. Bot. (ed. 2) 1: 632. 1840, nom. inval. Festuca procera Nees & Meyen, Nov. Actorum Acad. Caes. Leop.-Carol. German. Nat. Cur. 19(1): 166-167. 1843, nom. illeg. horn. Festuca orgyalis Willd. ex E. Fourn., Biol. Cent.Amer., Bot. 3: 582. 1885, nom. illeg. superfl. TYPE: Ecuador. Pichincha, Crescit locis alsis, subfrigidis regni Quitensos prope Chillo, San Antonio de Lulumbamba et Lloa, inter alt. 1280-1470 hexap., floret Aprili, Majo, Humboldt & Bonplands.n. (lectotype: B!, designated by Alexeev, Novosti Sist. Vyss. Rast. 23: 12. 1986; isolectotype: P!). 77 Festuca peruviana E.B. Alexeev, Bjull. Moskovsk. Obsc. Isp. Prir., Otd. Biol. 89(4): 114. 1984, horn, illeg. TYPE: Peru. Tabina, Jul 1854, Lechler 2114 (holotype: LE!; isotype: S). Festuca sublimis Pilg., Bot. Jahrb. Syst. 25(5): 718. 1898. TYPE: Ecuador. Azuay, Pucara de Chisalo, 1 Feb 1874,^. Stiibel 297 (holotype: B!; isotypes: BAA-1341 fragm. ex B!, US81580 ex B!). Tussocked perennials with intra- and extravaginal innovations. Culms 80-170 cm tall, erect, glabrous; nodes 1; cataphylls \-^ cm long, membranous to coriaceous, grayish-brown, striate. Leaf sheaths membranous to coriaceous, brownish-gray, glabrous or with short hairs; ligules 0.3-0.7 mm long, coriaceous, margins ciliate, apex truncate; blades 30-50 cm long, 0.8-1.6 mm wide, conduplicate, abaxially glabrous or sometimes scabrous, green, apex obtuse. Panicles 15-25 x 3•15(•20) cm, oblong to ovate, lax, densely flowered; branches glabrous or rarely sparingly scabrous. Spikelets 11-15 mm long, oblong-lanceolate, florets (4-)6 or 7; rachilla densely hairy; glumes (2-)3-5.5 mm long, membranous, glabrous below, upper 1/3 hairy, purplish-green, apex acute; lower glumes (2-)3-3.5 mm long, lanceolate, 1-nerved; upper glumes (4-)4.5-5.5 mm long, oblong, 3-nerved; lemmas (6-)6.5-8 mm long, lanceolate, 5-nerved, membranous to coriaceous, purplish-green, papillose, upper 1/3 scabrous, apex mucronate or shortawned, the awn 0.3-0.8 mm long; callus glabrous sometimes sparsely hairy; paleas as long as the lemma, lanceolate, membranous, hairy along upper 1/4 and keels, apex with longer hairs; lodicules ca. 1 mm long, obovate; anthers 2.8-3.5 mm long; ovary apex glabrous or with sparse hairs. Caryopses not observed. Leaf blade anatomy.•Cross-sections with (11-)15-21 vascular bundles and (5•)7•11 ribs above; sclerenchyma under abaxial epidermis continuous or discontinuous extending to all vascular bundle; adaxial epidermis discontinuous, extending to every second vascular bundle; adaxial epidermis with dense covering of curly to straight hairs, the hairs 0.15-0.2 mm long. Observations.•Festuca procera differs from other species of F. sect. Cataphyllophorae by forming large tussocks with tall culms (80-170 versus 60-80 cm tall in other members), long, densely flowered panicles, and long spikelets [(9-) 11-13 (-15) versus 6-11 (-12) mm in other members]. This 78 Festuca in South American Paramos Figure 36. Festuca procera. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 3313 (PRC). Festuca in South American Paramos -85 -80 -75 -70 79 -65 Figure 37. Distribution of Festuca procera (•), F. pilar-franceii (A), and Fboyacensis (•). species is morphologically quite variable. Specimens ofF. procera from southern Ecuador (region Azuay and Canar) present striking differences in abaxial surface texture of the leaf blades. Some specimens have sparingly scabrous leaf blades, inflorescence branches that are scabrous, and narrower panicles. In addition, all of these unusual specimens lack cataphylls, although this may be an artifact of the collection. Taxonomic status of this group of specimens needs further study. Distribution and habitat.•This species ranges from Colombia (Cordillera Central and Oriental), to Ecuador, and Peru. It occurs along margins of the Andean mountain forests and in matorral and transitional zones of grass paramo between 3400-3900 m. Additional specimens examined. COLOMBIA. Boyaca: Canon de Chicamocha, Mun. Sativasur, 17 Oct 1992, Etter 662 (COL); Mun. El Cocuy- Giiican, Parque Nacional El Cocuy, Hda. La Esperanza, Valle de los Frailejones, 3800 m, 30 Dec 1998, D. Stancik 1788 (COL, FMB, PRC). Cauca: Mun. San Sebastian, Macizo Colombiano, Valle de las Papas, 2910 m, 11 Sep 1958, Idrobo et al. 3737 (COL, US); Mun. Purace, Parque Nacional Purace, quebrada San Nicolas, 3100 m, 6 Apr 1985, Wood 4797 (COL, K). Norte de Santander: Quebrada del Rio Chitagu, 3400-3500 m, 20 Jul 1940, J. Cuatrecasas & Garcia-Barriga 10036 (COL); 3400-3500 m, 20 Jul 1940, J. Cuatrecasas & Garcia-Barriga 1003 7 (COL); Pamplona-Cerro alNE, 2770 m, 3400-3500 m, 26 Jul 1940, J. Cuatrecasas & Garcia-Barriga 10207 (COL). ECUADOR. Azuay: ca. 7 km S of road to Nabon, 03°07'S, 79°19'W, 3250-3350 m, S. Laegaard 101856 (AAU). Bolivar: km 14.5 E of Guaranda on road to Riobamba, 2950 m, P.M. Peterson 9297 & E.J. Judziewicz (K, MO, QCA, QCNE). Canar: km 11.6 80 Festuca in South American Paramos NW of Tambo on Panamerican Highway and 2.2 km E of road to microwave station, 3220 m, P.M. Peterson 8846, C.R. Annable & M.E. Poston (MO, QCA, US); S of El Tambo, ca. 1.5-3 km along road to Carshao, 02°28 'S, 78°59' W, 3100 m, S. Laegaard & Sklendf 20296 (AAU, LOJA, PRC); El Tambo, 3400-3500 m,D. Stancik 3793 (AAU, PRC, QCA); D. Stancik 3794 (PRC, QCA); Road Pimo-Santa Rosa, 3000 m, Acosta-Solis 21439A, B (US); Paramo road ca. 10 km WNW of Canar, 02°30'S, 79°00'W, 3550 m, S. Laegaard 71031 (AAU, PRC, QCA). Cotopaxi: at Laguna Quilotoa, 00°51'S, 78°53'W, 3500-3600 m, S. Laegaard 101350 (AAU, QCA); 3480-3500 m, S. Laegaard 101355 (AAU, QCA); 3700-3800 m, S. Laegaard 101340 (AAU, QCA); 00°52'S, 78°53'W, 3500-3750 m, Ceron 21862 (QAP); road Angamarca-Corazon km 11, 01°09'S, 78°57'W, 2880 m, 5 Apr 1992, S. Laegaard 102164 (AAU, QCA); Parque Nacional Cotopaxi, 3200 m, Izuriete 129 (AAU, QCA); 3500 m, Escalona & Gallegos 287 (US); Parque Nacional Cotopaxi, 2.4 km W of entrance, 2870 m, P.M. Peterson 8734, C.R. Annable & M.E. Poston (K, MO, QCA, QCNE, US); ca. 3 km de la entrada, 00°40'S, 78°30'W, 3200 m, 3 Jan 1982, Arguello 2, 33, 37, 440 (QCA); Sosa et al. 43 (QCA); 13 Nov 1982, Balslev & T. de Vries 3471 (QCA); km 2 N of Lasso on Panamerican Highway, 3270 m, 14 Apr 1990, P.M. Peterson 8747, C.R. Annable & M.E. Poston (K, MO, QCA, QCNE, US); Panam Highway, ca. 6 km N of Lasso, quebrada ENE Pastocalle, 3400 m, Sparre 15827, 15836 (S); Latacunga, paroquilla Sigchos, 2800 m, 30 Jun 1990, Ceron 10605 (QAP); Latacunga, paroquilla Belisario, Quevedo-Cerro Putzalagua, 00°57'S, 78°33'W, 2800-3500 m, Ceron 25977 (QAP). Chimborazo: road from San Fernando toward Mt. Chimborazo and El Arenal, 3930 m, Barclay & Juajiboy 8178 (COL, MO, US); Arenales de Palmira, ca. 7 km S of Guamote, 01°59'S, 78°43 'W, 3150 m, S. Laegaard et al. 18672 (AAU, LOJA, QCA,QCNE); ca. 7 km S of Palmira, 02°00'S, 78°43'W, 3170 m, 20 Mar 1992, S. Laegaard 101814 (AAU, QCA, QCNE); 02°02'S, 78°45'W, 3200-3300 m, S. Laegaard 71695 (AAU, QCA, QCNE); W of Arenales de Palmira, 02°01'S, 78°45 'W, 3330 m, S. Laegaard 105102 (AAU); km 8.9 N of Palmira, 3260 m, P.M. Peterson 8815 (K, MO, QCA, QCNE, US); km 10 on road AlaoChamlo, 01°50'S, 78°34'W, 3000 m, S Laegaard 55410 (AAU, QCA); km 4.6 of Pungala on road to Alao, 2970 m, P.M. Peterson 9179, E.J. Judziewicz, R.M. King & P.M. Jorgensen (K, MO, QCA, QCNE, US); km 9 S of Guamote, then 1.6 km SW on track to Alausi, 3300 m, P.M. Peterson 9316 & E.J. Judziewicz (K, MO, QCA, QCNE, US); Chimborazo, Chuquipoyo, 3900 m, Andre 3919 (K, US); ca. km 10 S of Palmira, 3250 m, 11 Nov 1985, S. Laegaard 55566 (AAU, QCA); ca. 5 km S of Palmira, 02°06'S, 78°45'W, 3300 m, S. Laegaard 20379 (AAU, LOJA); Mt. Chimborazo, Sodiros.n. (QPLS); Puela, 01°30'N, 78°30'W, 2315m, Ceron 15102 (QAP). Imbabura: Laguna Cuicocha, 00°18'N, 78°22'W, 3000-3100 m, S. Laegaard & S.A. Renvoize 70873, 70882 (AAU, K, QCA, QCNE); 2900-3100 m, 24 May 1991, Penafiel et al. 69 (MO, QCNE, PRC); 3100 m, E. Asplund 20176 (S); road Yahuarcocha-Mariano Acosta km 9,00°22'N, 78°03'W, 2900 m, 7 Feb 1992, S. Laegaard 101131 (AAU, QCA); Entre Muyurco y Tallachupa, Nevado Cotacachi, 3000 m, AcostaSolis 19006 (US); 3200-3300 m, Penafiel et al. 1187 (MO); Mun. Otavalo, road from Laguna Mojanda to Cochasqui, 00°04'55"N, 78°17'50"W, 3450 m, 19 Oct 2000, D. Stancik 4106 (PRC, QCA). Loja: km 1 E of Guachamana on road Macara-Catacocha, 2670 m, 9 Jun 1990, P.M. Peterson 9512 & E.J. Judziewicz (K, MO, QCA, QCNE, US). Morona-Santiago: Hda. HuarguallaHda. San Eduardo, 01°57'S, 78°32'W, 3600 m, 19 Jul 1999, D. Stancik 3312, 3313, 3314 (PRC, QCA); way to Parque Nacional Sangay, 02°15'S, 78°27'W, 3700 m, 19 Jul 1999, D. Stancik 3317 (PRC, QCA). Pichincha: road to Yanaurco on N side of Volcan Pichincha, 00°07'S, 78°34'W, 3600 m, 17 Jun 1981, S. Laegaard 52301 (AAU, QCA); Volcan Pichincha, 00°09'S, 78°32'W, 3200-3400 m, 5 Feb 1988, S Laegaard 70002 (AAU, QCA, OCNE); Sodiro s.n. (QPLS, US); km 41 on road Quito-Latacunga, 3350 m, 13 Apr 1990, P.M. Peterson 8717, C.R. Annable & M.E. Poston (QCNE, US); km 41, 3350 m, Buendia 2 (QCA); Environments of Quito, Jameson 191 (K); Panecillo, Sep 1887, Sodiro s.n. (QPLS); vicinity of Quito, 3100 m, E. Asplund 6151 (AAU, MO, S, US); Prop. Quito, Sodiro s.n. (QPLS, US); 3250 m, E. Asplund 6154 (AAU, S); 3500 m, 13 Jan 1926, E. Asplund 6139 (S, US); Road Pifo-Pintag, above Inga Monserrat 00°19'S, 78°17W, 34003500 m, S. Laegaard 102285 (AAU, QCA, QCNE); Sep 1890, Mt. Pichincha, Sodiro s.n. (MO, NY, QPLS); along road to the antennas, 00°09'S, 78°32'W, 3200-3400 m, S. Laegaard 70016 {AAU, QCA, QCNE); Mille 283, 285 (US); SE of Ungui, Festuca in South American Paramos 3000 m, Firmin 116 (US); above Quito, 3300 m, E. Asplund 16161 (S); 5 Feb 1988, S. Laegaard 70014 (AAU, QCA, QCNE); between Quito and Conoco to, 3000 m, E. Asplund 10375 (S); between Quito-Lloa, 3200 m, E. Asplund 7392 (S), 3600 m, Sparre 16006 (S); Volcan Cayambe, entrance to Parque Nacional, 00°03'S, 78°04'W, 3550 m, S. Laegaard & S.A Renvoize 70485 (AAU, K, QCA, QCNE); 3870 m, Escalona & GallegosE384 (MO); Antisana, road to Hacienda Pinatura, 3000-3100 m, 3 Mar 1988, S. Laegaard & S.A Renvoize 70542 (AAU, K, QCA, QCNE); 00°26'S, 78°20'W, 3400 m, S. Laegaard 102857 (AAU, QCA, QCNE). Tungurahua: road AmbatoGuaranda, km 25, 01°18'S, 78°48'W, 3300 m, 5 May 1999, S. Laegaard & Sanchez 20033 (AAU, LOJA, QCA); 01°24'S, 78°51'W, 4000^1100 m, S Laegaard 54802, 54809 (AAU, QCA); Pillaro, along irrigation ditch, 2850 m, 10 Aug 1939, E. Asplund 8135 (NY, QCA, S, US). Banos, 1800 m, 26 Sep 1923, A.S. Hitchcock 21926 (NY, US); Llangahua, 3300 m, Acosta-Solis 16678 (US); Pucara de Chisalo, Stiibel 297 (K). PERU. Ancash: Prov. Pallasca, 5 km S of road towards Cabana steep slope with Puya, Oxalis, Festuca, Tillandsia, 3210 m, 27 Mar 1997, P.M. Peterson 13922 & N. Refulio Rodriguez (K, MO, US, USM). Arequipa: Volcan de Misti, open slopes and canyon sides, 3100-3300 m, 4Nov 1925, Pennell 13231 (F). Cajamarca: Prov. Cajamarca, a 9 km de la Cajamarca, sobre carretera Cajamarca-Pacasmayo, 2800 m, 6 Nov 1984, Vega & Ruiz 3584 (MO); surrounding of Cajamarca, 2750 m, 21 May 1980, Becker et al. 1234 (LPB); Prov. Chota, near Las Palmas, ca. 24 km NE de Chota, fragmented mountain forest, 78°37'W, 06°29'S, 2789m, 17 Apr 1993, M. Dillon et al. 6359A (F, MO); Distr. Banos del Inca, a 3 km arriba de Banos del Inca, caretera Cajamarca-Celendin, 16 Mar 1984, Vega 3239 (F); km 13 de la carretera Cajamarca-Cumbe Mayo, frequent, 3000 m, 4 Sep 1984, Vega 3324 (F); Quebrada de la Esperanza, carratera a Cumbe Mayo, quebrada que converga al Valle de Cajamarca, arribe Cerro Sta. Apolonia, 2800 m, 4 Mar 1984, Vega et al. 3283 (F); Distr. San Juan, carretera Cajamarca-San Juan, 2350 m, 6 Dec 1993, Vega 723 (F); Distr. De Chetilla, ruta a Llullapuquio bosque perennifolio, 2650-2750 m, 21 May 1986, Vega et al. 4117 (F); Quebrada de la Esperanza, carratera a Cumbe Mayo, quebrada que converga al Valle de Cajamarca, arribe Cerro Sta. Apolonia, 2800 m, 4 Mar 1984, Vega et al. 3283 (F); a 1 km n arriba de San Juan, siguiendo la carretera a Cajamarca, 2300 m, 6 Feb 1985, Vega et al. 3855 (F); Sexcemayo, al W de la ciudad de Cajamarca, 3500 m,20Jun 1991, Vegaetal. 5745 (F); a la altura del Paso El Gavilan, 3200 m, 18 Apr 1976, Vega et al. 1393 (F); Entre La Encanada y Kumullca, 3300 m, 17 Jun 1975, Vega et al. 1598 (F); Prov. Celendin, Guanambra, a 11 km de la carretera Celendin-Cajamarca, 2830 m, 3 Dec 1988, Vega 4661 (F); al E de la ciudad de Celendin, 2600 m, 5 Apr 1980, Vega 3830 (F); Celendin-Cajamarca road, 78°12'W, 06°58'S, 2800-3400 m, 25 Feb 1984, Smith 6222 (MO); above Celendin, 2750 m, 23 Mar 1988, S.A. Renvoize 4879 (K); Bajando el paso de Gelig, sobre la carretera Celendin-Balsas, 3000 m, 15 Jun 1981, Vega 2576 (F); Prov. Chota, 5 km N of Chota on road towards Conchan, 2560 m, 18 Mar 2000, P. M. Peterson 14968 & N. Refulio Rodriguez (MO, US, USM); 20 km NW of Bambamarca on Hwy 3 N towards Chota, 2900 m, 18 Mar 2000, P.M. Peterson 14953,14955,14957 &N. Refulio Rodriguez (K, MO, US, USM); Chota-Tacabamba road, 6-8 km from Chota, 78°38'W, 06°32'S, 2650 m, 20 Feb 1983, Smith et al. 3627 (K, MO); Distr. Llacanora, sobre la carretera a Cajabamba, 2760 m, 6 Jul 1980, Vega 2250 (F); Prov. Cutervo, 10 km NW of Chiguirip and 19 km SE of Cutervo, 2630 m, 19 Mar 2000, P.M. Peterson 14989,14992 & N. Refulio Rodriguez (K, MO, US, USM); Prov. Hualgayoc, cerro La Llama de las Ventanillas, al SW de Bambamarca, 3 890 m, 5 Mar 1991, Vega et al. 5696 (F); Prov. San Marcos, Yanupacha, al E de Ichocan, 3100 m, 26 Mar 1994, Vega et al. 6933 (F). Cuzco: Prov. Paucartambo, altura de Teleban, 3700-3800 m, 16 Jul 1990, Cano3780 (F); Huayna Pichu, rocks, 68°07'W, 16°17'S, 3000 m, 5 Aug 1937, Tutin 1285 (BM). Junin: Huancayo, 18.5 km NW of Huancayo up Rio Shullcas, just above Acopalca, 3890 m, 12 Apr 1997, P.M. Peterson 14224 & O. Tovar(K). La Libertad: Prov. Bolivar, arriba de Longotea, ruta Bolivar, 2800 m, 28 Aug 1989, Vega 5013 (F). Lambayeque: Prov. Ferrenafe, Incahuasi, 3400 m, 22 Jun 1986, Quiroz 1929 (F, MO). Piura. Prov. Huancabamba, San Antonio, km 25 de la carretera Huancabamba-Salala, 2850 m, 5 Jan 1990, Vega etal. 5185 (F); 30 kmN of Huancabamba and 3 km N of Salala, 29803200 m, 2 Apr 2000, P.M. Peterson 15178 & N. Refulio Rodriguez (K, US, USM). 30. Festuca toca Stancik, Darwiniana41(l^l): 123, f. lOh-m. 2003. (Figs. 38,39,88A & B). TYPE: 82 Festuca in South American Paramos 2 mm 1 mm Figure 38. Festuca toca. A. Habit. B. Inflorescence. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 1418 (PRC). Festuca in South American Paramos Colombia. Boyaca, Mun. Toca, road from Pesca to Toca, km 4-5, Paramo Cortadero, 5°30'N, 73°15'W,2700m, 14Nov 1998,/). Stancikl404 (holotype: PRC!; isotypes: COL!, FMB!). Rhizomatous perennials with extravaginal innovations. Culms 50-90 cm tall, erect, solitary, glabrous or finely scabrous; nodes 2-A; cataphylls short, coriaceous, gray-brown. Leaf sheaths membranous, brown, striate; auricles absent; ligules 0.10.5 mm long, membranous, apex truncate; blades 10-35 cm long, 0.5-1.1 mm wide, conduplicate or conduplicate to involute, green, abaxially glabrous. Panicles 6-15(-20) x ca. 0.5 cm, erect, contracted, elongate; branches scabrous. Spikelets 6-8 mm long, florets 4 or 5(-6); rachilla glabrous or with scattered hairs; glumes 2•4 mm long, lanceolate, membranous to coriaceous, purplish, apex acute, scabrous; lower glumes 2-3 mm long, 1-nerved; upper glumes 3-4 mm long, 3-nerved; lemmas -80 -75 83 4-5 mm long, lanceolate, 5-nerved, membranous to coriaceous, purplish, upper 1/3 scabrous, apex mucronate; callus glabrous; paleas as long as the lemma, membranous, papillose, upper 1/3 hairy, lodicules lanceolate, acuminate; anthers 1.1-1.6 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 3/5 of total lenght. Leaf blade anatomy.•Cross-sections with 7-15 vascular bundles and 5-13 ribs above; sclerenchyma under abaxial epidermis discontinuous, sometimes abaxial continuous, extending to some (rarely to all) vascular bundles forming girders; adaxial epidermis with scattered hairs, the hairs 0.2-0.9 mm long. Observations.•Festuca toca differs from the remaining species of F. sect. Cataphyllophorae by having solitary culms. Festuca toca is morphologically similar to F. andicola (F sect. Aulaxyper), but differs by having longer glumes (lower glumes 2-3 versus 1.4-1.8 mm; upper glumes 3-4 versus -70 Figure 39. Distribution of Festuca asplundii (•), F. toca (A), and F. cocuyana (•). -65 84 Festuca in South American Paramos 2-2.5 mm) and longer anthers (1.1-1.6 versus 0.8-1 mm). Distribution and habitat.•This species is endemic to Colombia where it is known only from the Cordillera Oriental (Boyaca, Norte de Santander). It occurs in swampy patches in grass paramo zone between 3100-3500 m. Festuca toca is present in different vegetation communities such as: Lysipomia sphagnophilla subsp. minor (Cleef 1981). Additional specimens examined. COLOMBIA. Boyaca: Mun. Duitama, paramo Pan de Azucar, 3450 m, 30 Nov 1998, D. Stancik 1418 (COL, PRC); Paramo de Rusia, 3400 m, 2 Jan 1984, Wood 4481 (COL, FMB, K); 3570 m, 13 Dec 1972, Cleef 7139 (COL); summit of road from Duitama to Abendanos, 3490m, 12Feb 1972,/). Stancik2371 (COL, PRC); Hda Los Tres Corrales, 5°55'3.2"N, 73°4'13.5"W, 3490 m, Hernandez 1022A (COL); Mun. Toca, road from Pesca to Toca, km 4-5, paramo Cortadero, 5°30'N, 73°15'W, 2700 m, 14Nov 1998,D. Stancik 1404(COL,FMB,?RC); 3350 m, 14 Nov 1998, D. Stancik 1365 (COL, FMB, PRC); Mun. Susacon, road km 20, margin of Rio Susacon, 3150 m, 24 Feb 1999, D. Stancik 2513 (COL, PRC); Paramo Guantiva (Sanjuanero), 3300 m, lOAug 1977, Ruiz & Romero 3 (COL); Mun. Paipa, Cuchilla El Paramo, 3200 m, 4 Dec 1998, D. Stancik 1533 (COL, PRC); Mun. Arcabuco, Santuario Iguaque, way from Laguna Iguque to Laguna Ojo de Agua, 3650 m, 15 Oct 1998, D. Stancik 948 (COL, FMB, PRC). Norte de Santander: Paramo de Santurban, 3200 m, 4 Jan 1984, Wood 4507 (COL, FMB, K). 31. Festuca asplundii E.B. Alexeev, Bjull. Moskovsk. Obsc. Isp. Prir, Otd. Biol. 89(4): 116. 1984. (Figs. 39, 40, 88C-F). TYPE: Ecuador. Pichincha, Iter Regnellianum quartum, Flora Aequatoriensis, prov. Pichincha, vicinity of Quito, Rucu Pichincha, 4650 m, 31 Aug 1939, E. Asplund 8586 (holotype: S!). Tussocked perennials with intravaginal innovations. Culms up to 80 cm tall, erect, scabrous; nodes 1 near base. Leaf sheaths wide, coriaceous, stramineous, scabrous; ligules 0.5-1.5 mm long, coriaceous, apex truncate or emarginate, short-ciliate; blades ca. 40 cm long, 1-1.4 mm wide, conduplicate to involute, rigid, somewhat abaxially scabrous, green, apex obtuse. Panicles 12-27 x 3^4(•10) cm, contracted, rarely with open branches; branches scabrous. Spikelets sterile, forming l(-2) shoots (i.e., vegetative proliferation), florets 1•4, up to 6 when shoots lacking; glumes 4.5-6(-7) mm long, coriaceous, dark purple, scabrous dorsally, distal margins (upper 1/2) membranous, apex acute; lower glumes 4.5-5(-6) mm long, lanceolate, 1-nerved; upper glumes 5-6(-7) mm long, oblong-lanceolate, 3-nerved; lower lemmas sterile, 4.5-6(-7) mm long, 5-nerved (sometimes inconspicuously), lanceolate, scabrous or papillose dorsally, margins membranous, apex entire or slightly two-dentate and short-awned, the awns up to 0.8 mm long, dark purple; lodicules and sexual organs absent; upper lemmas converted into prolifering shoots 1-2.5 cm long, 1-3-leafed are prolonged into leafy bracts. Leaf blade anatomy.•Cross-sections with 11-17 vascular bundles and 9-15 ribs; sclerenchyma under abaxial epidermis continuous extending to all the vascular bundles; adaxial epidermis discontinuous extending to every other vascular bundle; bulliform cells absent; abaxial epidermis with prickles and densely hairy adaxially, the hairs 0.04-0.09 mm long. Observations.•This species does not seem to be morphologically similar to any other species of Festuca from South America. St.-Yves (1927) considered F. asplundii to be a viviparous form of F procera only lacking cataphylls. Individual specimens of F asplundii are highly variable in spikelet size, panicle shape, and ligule characteristics. We include numerous specimens from Peru in F asplundii, although Alexeev considered this species to be endemic only to Colombia and Ecuador. He attributes Peruvian specimens to the unclearly described species, F. ancachsana and invalidly described, F peruviana (Alexeev 1986). In Festuca, vivipary occurs rarely in developing spikelets, although, it can sometimes be found in a few individuals of F subulifolia, F procera, and F tolucensis. Distribution and habitat.•Festuca asplundii is known from northern Peru, Ecuador, and Colombia (Cordillera Central and Cordillera Occidental). It is found in swampy and humid patches of grass paramo and the lower zones of superparamo between 3300-4300 m. It is a dominant and co-dominant species in communities of Espeletia hartwegiana subsp. centroandinay Calamagrostis recta (Cleef et al. 1983), Festuca dolichophylla, Senecio latiflorus & Pentacalia vernicosa (Sturm and Rangel 1985), Festuco dolichophyllae-Calamagrostietum effusae Festuca in South American Paramos 85 2 mm Figure 40. Festuca asplundii. A. Stylized growth form. B. Habit. C. Inflorescence. D. Ligule. E. Spikelet. F. Glumes. G. Lemmas. H. Leaf blade cross-section. A-H, Stancik 3675 (PRC). 86 Festuca in South American Paramos (Salamanca et al. 1991), Senecionetum rufescentisAgrostiosum (Cuatrecasas 1934), Calamagrostis effusa and Loricaria cf. complanata (Rangel et al. 1995), Espeletia pycnophylla & Arcytophyllum capitatum (Rangel and Luteyn 1995), GynoxyoPolylepidetum sericae (Salamanca et al. 1991), Loricaria thuyoides & Pentacalia vernicosa (Rangel et al. 1995), Loricaria thuyoides & Arcytophyllum capitatum (Rangel et al. 1995), Calamagrostis effusa & Festuca aff. dolichophylla (Salamanca 1991), and Calamagrostis recta, Festuca aff. sublimis & Diplostephium rupestre (Cuatrecasas 1934). Additional specimens examined. COLOMBIA. Caldas: Mun. Manizales, Parque Nacional Los Nevados, entrance, 4020 m, 18 Sep 1999, D. Stancik 3410, 3411 (COL, PRC); road from Casa del Cisne to Rio Nereidas, km 5, 3800^000 m, 18 Sep 1999, D. Stancik 3401, 3402 (COL, PRC); Carretera Manizales-Nevado del Ruiz, km 72,4100 m, 1 Mar 1977, Forrero et al. 3644 (COL, MO); Carretera entre Manizales y el Nevado del Ruiz, 4000 m, 5 May 1959, Pinto-Escobar 415 (COL, PSO); Paramo de Herves, division de Aguato, 3300 m, Feb 1852, Triana 810 (COL); 3500 m, Feb 1852, Triana s.n. (US); alrededores del Nevado del Ruiz, 4140^300 m, 9 Oct 1978, Rangel et al. 1800 (COL); Nevado del Ruiz, 8 Oct 1983, Wood 4030 (AAU, K); Paramo El Ruiz, 3700-4200 m, 26 Dec 1936, Chardon 5010 (COL); 3400-3700 m, 17 Dec 1917, Pennell 3064 (K, MO, US); Nevado del Ruiz, arenales de Ruiz, 3 km a de la Olleta, 4250 m, 14 Oct 1972, Cleefet al. 5983 (COL, U, US); 3900-4200 m, 5 May 1940, J. Cuatrecasas 9311 (COL, US); Carretera El Ruiz-Otun, Hda. Buenos Aires, 3940 m, 20 May 1979, Salamanca AC'172(COL); 3500m, Dec l974,Llana 71 (MEDEL); Paramo de Quindio, 4300-4500 m, 20 Aug 1922, Pennell 9890 (K, US). Cauca: Mun. Popayan, Parque Nacional Purace, Pilimbala, 02°20.07N, 76°23.65 'W, 4200^300 m, 6 Jul 2000, D. Stancik 3611 (COL, PRC); Purace, Laguna San Rafael, 3300 m, 6 Apr 1985, Wood 4802 (COL, FMB, K). Nariflo: Mun. Cumbal, Nevado del Cumbal, N-NE slope, 4100 m, 9 Mar 1999, D. Stancik 2724 (COL, PRC); Mun. Tuquerres, Volcan Azufral, Laguna Verde, 3750 m, 9 Mar 1999, D. Stancik 2759 (COL, PRC); Mun. Pasto, Volcan Galeras, 3920 m, 6 Aug 1977, Pinto-Escobar 1845 (COL); 3900 m, 29 Oct 1983, Wood 4062 (COL); Volcan Galeras, 3900 m, Wood 4063 (FMB, K). Risaralda: Mun. Santuario, SE del Cerro Venta- nas, Macizo del Tamara, 3820 m, 11 Feb 1983, Torres-Romero et al. 1875 (COL); Mun. Pereira, Parque Nacional Los Nevados, Laguna del Otun, 3800 m, 16 Feb 1980, Jaramillo-Mejia et al. 5634 (COL); Nevado de Sabanta Isabel, 4300 m, 17 Feb 1980, Jaramillo-Mejia et al. 5683 (COL). Mun. Santa Rosa, Volcan Santa Rosa, 4280 m, 20 Feb 1980, Jaramillo-Mejia et al. 5770 (COL); Paramo entre Termales y Libano, 4000 m, 21 Dec 1958, Barclay & Juajibioy 6456 (COL, MO, US). Tolima: Mun. Ibague, 04°38.5N, 75°19.1 'W, 4250 m, 8 Jun 2000, D. Stancik 3606 (COL, PRC); Mun. Santa Isabel, vertiente oriental abajo del paso de El Otun, 4200 m, 7 Feb 1980, Jaramillo-Mejia 623 7 (COL); Quebrada Africa, 4200^300 m, 5 Feb 1980, DiazPiedrahita & Jaramillo-Mejia 1821 (COL, U). Valle de Cauca: Mun. Cali, Farallones de Cali, 03°19.78N, 76°41.7'W, 3950 m, 9 Jul 2000 D. Stancik3612, 3613, 3614 (COL, PRC); 3650-3700 m, D. Stancik 3637 (COL, PRC). ECUADOR. Azuay: Parque Nacional Cajas, TotorocochaMazan Valley, 02°53'S, 79°10'W, 4000 m, 12 Nov 1987, Ramsay & Smith 502 (K, QCA, QCNE); Cuenca-Molleturo road, 3600-4200 m, 26 Jul 1982, Clements et al. 2170 (AAU, QCA, QCNE); Paramo de Soldados ca. 14 km above Soldados, 02°08'54"S, 79°08'17"W, 3900 m, 9 Jan 2000, S Laegaard et al. 20931 (AAU, LOJA); between Huagrarancha and Loma de Galapagos, 3140-3505m, J. Steyermark 53481 (US); Mun. Cuenca, Parque Nacional Cajas, N side of Lagoon Taglococha, 02°47'S, 79°15'W, 4100^1200 m, 1 Sep 2000, D. Stancik 3849, 3871 (PRC, QCA). Carchi: Volcan Los Chiles, along road ca. 1 km E of pass, 00°49N, 77°57'W, 3950-3980 m, S. Laegaard 101695 (AAU, QCNE); D. Stancik 3241, 3279 (PRC, QCA); 00°48'N, 77°56'W, 3850^000 m, S. Laegaard 54972 (AAU, QCNE); SW slope of Volcan Chiles, 3740 m, B. Ollgaard & Balslev 8545 (AAU); S slope, Ramsay 896 (K, QCA); Paramo del Angel, 3700 m, S Largaard 101284 (AAU, QCA); sector El Volador, 00°38'N, 67°53'W, 3400-3800 m, Davalos 19 (US). Chimborazo: E slope of Mt. Chimborazo, 4450 m, E. Asplund 7927 (S); 4250 m, E. Asplund 7806 (S); El Altar, N side of the Volcan, 4200^300 m, Sklendf & Kosteckovd 1106 (QCNE); Paramo de los Altares, Collanes valley, 4200 m, Ramsay & Smith 415B (K, QCA, QCNE); N side of Volcan, 4200^400 m, Sklendf & Kosteckovd 1028 (AAU); 4300 m, Sklendf & Kosteckovd 97-10 (QCA); N side, 4400 m, Sklendf Festuca in South American Paramos & Kosteckovd 91-8 (US); from campsite above Rio Alao 8.5 km E of Guardiana Alao, 3350-3550 m, P.M. Peterson 9204, E.J. Judziewicz, R.M. King & P.M. Jorgensen (K, MO, QCA, QCNE, US); W of pass Alao-Huamboya, 01°48'S, 78°25'W, 37503800 m, S. Laegaard 55425 (AAU, QCA). Cerro Yanaurcu, 02°14'S, 78°30'W, 4000-4200 m, Sklendf & Kosteckovd 1478 (AAU); Paramo de Las Tres Cruces, N and E of Alao, 3500 m, Barclay & Juajibioy 8804 (MO, US); between Mt. Chimborazo and Urbina, 3600-4500 m, A.S. Hitchcock 21978 (US). Cotopaxi: Cerro Verde, road San Miguel-Puerto Nuevo, km 29, 3950^1050 m, B. Ollgaard & Balslev 9924 (MO). Imbabura: Proantag, 3500-3800 m, Acosta-Solis 19199 (US); Mun. Urcuqui, road to Cerro Yanaurcu, 00°28.5 'N, 78°20'W, 4150 m, 15 Oct 2000, D. Stancik 4097 (PRC, QCA). Mun. Cayambe, Volcan Cayambe, 00°00'31.6"N,78°00'55.6"W,4450m,20Oct2000, D. Stancik 4150, 4151 (PRC, QCA). Loja: Jionbura-Yumba road, ca. 3 km W of pass, 04°44'S, 79°25'W, 3450 m, S. Laegaard et al. 18606 (AAU, LOJA, QCA, QCNE); along road to Fierra Urcu, ca. 10 km from main road Loja-Saraguro, 03°33 'S, 79° 15' W, 3400 m, S. Laegaard et al. 18859 (AAU, LOJA, QCA, QCNE); Cordillera de Amaluza, Laguna de Jimbira, 04°42'S, 79°25'W, 3400 m, Cabrera 438 (LOJA); Laguna Chuquiraga E of Amaluza, 04°37'S, 79°22'W, 3300 m, S. Laegaard etal. 19280 (AAU, LOJA, QCNE); Mun. Saraguro, road to Fierra Urcu, 03°42'40"S, 79°18'12"W, 3400-3450 m, 24 Aug 2000, D. Stancik 3 763,3 767 (PRC, QCA). Napo: Parque Nacional Las Llanganatis, base of Cerro Hermoso on W side, near lake 78°18'W, 01°14'S, 3850 m, 13 Nov 1999, D.A. Neill et al. 12042 (MO); W of Cerro Hermoso, near saddle between headwaters of Rio Verde and Rio Topo, 78°19.5'W, 01°11'S, 3950 m, D.A. Neill et al. 11992 (MO); camino desde el Paramo de Soguillas hasta Aucacocha, 78° 19'W, 01 °08 'S, 3940 m, 16 Nov 1999, Narvazez et al. 548 (MO); Paramo de Soguillas, near Las Torres de Llanganatis, 3 850^000 m, 16 May 1982,5. Ollgaard & HolmNielsen 38728 (MO). Pichincha: W side of Sincholagua, 13-14,500 ft, Feb 1880, Whymper 1326 (K). W side of the mountain ridge ca. 2 km W of Cerro Saraurcu, 4200 m, Sklendf & Kosteckovd 1817 (AAU); Nevado Cayambe, W side, 4200 m, Sklendf & Kosteckovd 1142 (QCA); SW slope, 4400 m, Sklendf & Kosteckovd 1890 (AAU); SW slope, 4250 m, Molau & Eriksen 3244 (QCA); NE 87 side of Cayambe Mountain, ca. 4900 m, Cazalet & Pennington 5763 (K, US); 3 km S of Refugio on the SW slope, 4220 m, P.M. Peterson 9076, E.J. Judziewicz & R.M. King (K, MO, QCA, QCNE, US); along road to refuge, 00°04'S, 77°57'W, 4300 m, S. Laegaard & S.A. Renvoize 70511 (AAU, K, QCA, QCNE); W side, 4200 m, Sklendf & Kosteckovd 64-19 (US); N slope, 3750-3800 m, B. Ollgaard et al. 34202 (K, MO); base of Volcan Sincholagua, 00°34'S, 78°22'W, 4200 m, HolmNielsen 6612 (AAU, S); Cerro Sincholagua, 3900-4000 m, Balslev et al. 3901 (AAU, MO, QCA); W side, 13,000-14,500 ft, Whymper s.n. (K); Paramo de Guamani, E. Asplund 17174 (S); Laguna Linda, 3 km W of Paso de Guamani, 3900 m, Sparre 17719 (S); 01°19'S, 78°12'W, 4000 m, S. Laegaard 18449 (AAU, LOJA, QCA, QCNE); 00°56'S, 78°23'W, 3600 m, S Laegaard 53343 (AAU, QCA, QCNE); 4050 m, B. Ollgaard & Balslev 10122 (MO); Laguna de Papallacta, 3750 m, B. Ollgaard & Balslev 8170 (MO); antenna, 4000-4100 m, B. Ollgaard & Balslev 10074 (M); Aguirre et al. 4206 (LOJA); 4050 m, S. Laegaard 51318 (AAU, QCA); S of Paso de la Virgen, 00°21'S, 78°13'W, 4050^250 m, S Laegaard 55680 (AAU, QCA); 4200-4250 m, S. Laegaard 101383 (AAU, QCA); W slopes, 38004100 m, Balslev 1615 (AAU); 4060 m, Fierrp 24 (QCA); 4050 m, Naranyo 4 (QCA); 3960 m, Len 1139 (QCA); Volcan Cotacachi, 00°35 'S, 78°20'W, 4150 m, Ramsay & Smith 785 (K, QCNE); 4200 m, Sklendf & Kosteckovd 112-25 (QCA); vicinity of Quito, Rucu Pichincha, 465 m, E. Asplund 8576 (MO, NY, QCA, S); Volcan Antisana-Laguna Micacocha, B. Ollgaard & Balslev 8868 (MO); 3850-3950 m, S Laegaard 101595 (AAU, QCA, QCNE); valley NW of N peak of Volcan, 40004300 m, Grubb et al. 639 (K); Paramo de Mojanda, between Laguna Grande and L. Negra, 00°08N, 78°16'W, 3700-3800 m, S. Laegaard 54581 (AAU, QCA, QCNE); Mun. Pifo, Paramo de Guamani, grass paramo, 00°19'S, 78°12'W, 4300 m, 19 Jun 1999, D. Stancik 3044, 3050 (PRC, QCA); Mun. Amaguana, summit of Volcan Pasachoa, 00°28'S, 78°29'W, 4200 m, 14 Sep 2000, D. Stancik 3675 (PRC, QCA, US); 4150 m, D. Stancik3703 (PRC, QCA). Sucumbios: Paramo Mirador, SW of Playon de San Francisco, 3400-3600 m, P.M. Peterson 9154, E.J. Judziewicz & R.M. King (US). Tungurahua: Cordillera de Llanganatis, El Tambo near Lake Yanacocha, 3650 m, E. Asplund 10000 Festuca in South American Paramos (S); below summit of Pan de Azucar, 4100 m, B. Ollgaard et al. 38541 (AAU, MO, NY, QCA); B. Ollgaard et al. 38540 (AAU); N slope towards Rio Golpe just N of Chosca Aucacocha, 3600 m, B. 0llgaard et al. 38670 (AAU, MO, NY, QCA); Las Tolas, ca. 5 km SE of Laguna Pisayambo, 0P07'S, 78°21 'W, 3900^1000 m, S. Laegaard et al. 20733 (AAU, LOJA); Paramo de Jaramillo, 01°08'10"S, 78°08'22"W, 4000-4250 m, S. Laegaard 53289 (AAU, QCNE); Faldas de Cazahuala, 3200 m, Acosta-Solis 5097 (US); Mun. Pillaro, Las Llanganatis, Rio Milin, 01°07'37"S, 78°21'W, 3750 m, 28 Sep 2000, D. Stancik 3904 (PRC, QCA). Zamora-Chinchipe: Sendero AmaluzaPalanda, cerca de la Laguna Arrebatadas, 3350 m, Larsen et al. 21 (AAU, NY, QCA). PERU. Amazonas: Prov. Chachapoyas, upper slope of Cerro Campanario, 3600-3900 m, 3 Aug 1962, J. Wurdack 1572 (US). Ancash: Yungay Prov., Huascaran Parque Nacional Llanganica Sector, Quebrada Ancachs at portachuelo, 09°03 'S, 77°35 'W, 47704870 m, 31 Dec 1984, Smith et al. 8879 (QCA, U); Yungay Prov., Huascaran Parque Nacional, 4700 m, Smith 11318 (MO, QCNE); Prov. Huari, CatacChavin road, 40 km from Catac, Huascaran Parque Nacional, 9°42'S,77°13'W, 4360 m, 18Aug 1984, Smith 8284 (MO, US); Prov. Recuy, Huascaran Parque Nacional, camino over Cahuish Tunnel, puna, 09°41'S,77°15'W, 4550-4500 m, 7 Oct 1985, Smith et al. 11127 (MO); Prov. Yungay, Huascaran Parque Nacional, Morococha at largest lake 08°55'S, 77°35'W, 4550 m, 15 Jan 1985, Smith et al. 9225 (MO). Cajamarca. Prov. Cajamarca. 52 km N of Cajamarca on Hwy 3 N towards Bambamarca, and ca. 1 km W on a small road, Carretera Lagunas Yanacanchilla, 3800 m, 16 Mar 2000, P.M. Peterson 14892 & N. Refulio Rodriguez (MO, US, USM); Prov. Hualgayoc, desvio de la carretera Coymolache-Chugur, Cerro Tantahuatay, 3700 m, 29 Apr 1994, Vega 7087 (F); CajamarcaBambamarca road, 55 km N of Cajamarca, 06°55'S, 78°35'W, 3750 m, 6 Feb 1984, Smith et al. 7428 (MO). La Libertad: Prov. Bolivar, entre Longotea y Bolivar, en jalca, 3600 m, 2 Jun 1960, Lopez & Sagasteguii 3340 (US); Prov. Pataz, Paso de Alaska, Carretera a Tayabamba, 3900 m, 24 Jun 1974, Lopez & Sagasteguii 8180 (MO). San Martin. Prov. Mariscal Caceres, grassland in NW sector Rio Abiseo Parque Nacional, 07°40'S, 77°30' W, 3650 m, B. Leon & Young 1662 (MO). 32. Festuca boyacensis Stancik, Darwiniana 41(1^1): 128, f. 7f-k. 2003. (Figs. 37,41,89A & B). TYPE: Colombia. Boyaca, Mun. Aquitania, paramo Los Curies, 3500 m, 8 Feb 1999, D. Stancik & Medina 2166 (holotype: PRC!; isotypes: COL!, FMB!). Tussocked perennials with intravaginal innovations. Culms 30-60 cm tall, erect, glabrous; nodes 1 near base with 2 short leaves. Leaf sheaths membranous, stramineous, occasionally brownish, glabrous; auricles absent; ligules 0.3-0.6 mm long, membranous, apex truncate, ciliate; blades 15-20 cm long, 0.4-0.7 mm wide, conduplicate to involute, glaucous, abaxially finely scabrous, apex acute. Panicles 6-15 x 0.5-1 cm, contracted, narrow; branches scabrous. Spikelet 8-9.5 mm long, lanceolate, florets 3 or 4; rachilla glabrous; glumes 3-5.2 mm long, membranous, glabrous, margins finely ciliate, purplish, apex acute; lower glumes 3-3.7 mm long, lanceolate, 1 -nerved; upper glumes 4-5 mm long, ovate, 3-nerved; lemmas 6-7.5 mm long, lanceolate, 5-nerved, membranous, glabrous, purplish, awned, the awn 1-2 mm long; callus glabrous; paleas 5/6 as long as the lemma, glabrous, keels ciliate; lodicules lanceolate, acuminate; anthers 0.9-1.2 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 1/2 as long as the grain. Leafblade anatomy.•Cross-sections with 5-7 vascular bundles and 3-5 ribs above; sclerenchyma only abaxially present, discontinuous or almost continuous; bulliform cells absent; adaxial epidermis with sporadic hairs, the hairs 0.01-0.1 mm. Observations.•Festuca boyacensis is morphologically similar to the Ecuadorian species, F. chimborazensis. However, F. chimborazensis has smaller culms [8•15(•28) versus 30-60 cm], smaller panicles (2-8 versus 6-15 cm), and glumes with obtuse (acute in F. boyacensis) apices. Distribution and habitat.•Festuca boyacensis is endemic to Colombia, known only from the Cordillera Oriental (Boyaca). It occurs in swampy patches and margins of lakes and springs in grass paramos between 3000-3500 m. Additional specimens examined. COLOMBIA. Boyaca: Mun. Santa Rosa de Viterbo, paramo Alto Lamadero, 3200 m, 30 Nov 1999, D. Stancik 1493 (COL, MB, PRC). 33. Festuca carchiense Stancik, Folia Geobot. Phytotax. 39(1): 98, f. 1, 1-5. 2004. (Figs. 42, 44, Festuca in South American Paramos 89 Figure 41. Festuca boyacensis. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 1493 (PRC). 90 Festuca in South American Paramos Figure 42. Festuca carchien.se. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Laegaard 101162 (AAU). Festuca in South American Paramos 89C & D). TYPE: Ecuador. Carchi, km 11 along road Las Juntas (Tulcan)-El Angel, 77°50'W, 00°43'N, 3330 m, 11 Mar 1992, S. Laegaard 101716 (holotype: AAU!; isotypes: PRC!, QCA!, QCNE!). Densely tussocked perennials with intravaginal innovations. Culms 50-60 cm tall, erect, glabrous; nodes 1, basal. Leaf sheaths membranous, brownish-gray, glabrous, inconspicuously striate; ligules 0.5-0.8 mm long, apex short-ciliate; blades 25^10 cm long, 0.4-0.7 mm wide, conduplicate to involute, finely abaxially scabrous, green, apex obtuse. Panicles 10-20 x ca. 0.7 cm, contracted, slender; branches scabrous. Spikelets 9-10.5 mm long, lanceolate, florets 3 or 4; rachilla with scattered hairs; glumes (3.5-)4-6.5 mm long, membranous, purplish-green, upper 1/5 scabrous; lower glumes (3.5-)4-4.5 mm long, oblong-lanceolate, 1-nerved, apex acute; upper glumes 5-6.5 mm long, oblong, 3-nerved, apex acute; lemmas (6-)7-7.5 mm long, lanceolate, 5-nerved, membranous, purplish-green, scabrous distally, awned, the awn 0.8-1.5 mm long; callus with scattered hairs; paleas as long as the lemma, membranous, upper 1/3 and keels scabrous to hairy; lodicules ca. 0.7 mm long, lanceolate; anthers 1.1-1.3 mm long; ovary apex glabrous. Caryopses not observed. Leaf blade anatomy.•Cross-sections with 5 vascular bundles, 3(-5) ribs above; sclerenchyma discontinuous and small under abaxial epidermis, absent under adaxial epidermis; adaxial epidermis with scattered hairs, the hairs ca. 0.09 mm long. Observations.•Festuca carchiense is morphologically similar to F. glumosa and F imaburensis. (See details in F glumosa section). Distribution and habitat.•This species is known only from northern Ecuador (Carchi, Imbabura) and it occurs in grass paramo between 3200-3800m. Additional specimens examined. ECUADOR. Carchi: Paramo del Angel, lakes at Los Voladeros, 00°41'N, 77°53'W, 3700 m, S. Laegaard 101294 (AAU, QCA); 3200-3800 m, S. Laegaard 55769 (AAU). Imbabura: road Yahuarcocha-Mariano Acosta, km 20, Paramo de Mariano Acosta, 00°20'S, 78°00'W, 3600-3650 m, 8 Feb 1992, S. Laegaard 101162 (AAU, QCA, PRC). 34. Festuca chimborazensis E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 69: 1549. 91 1984. (Figs. 43,44,90A & B). TYPE: Ecuador. Chimborazo, Iter Regnellianum quartum, Flora Aequatoriensis, prov. Chimborazo, southern slopes of Mt. Chimborazo, hard dry ground along a rivulet, 3900 m, 21 Aug 1939, E. Asplund 8446 (holotype: S!; isotypes: NY!, QCA!). Densely tufted perennials with intravaginal innovations. Culms 8-15(30) cm tall, erect, glabrous; nodes 1, basal with a single short leaf. Leaf sheaths coriaceous, grayish, finely striate, glabrous; ligules 0.3-0.4 mm long, membranous, apex truncate, short-ciliate; blades 4-15 cm long, 0.4-0.6 mm wide, conduplicate to involute, curved to mostly straight, glaucous, apex obtuse. Panicles 2-10 cm long, 5-7 mm wide, contracted, narrow; branches scabrous on all surfaces. Spikelets 8•10(•11.5) mm long, lanceolate, florets 3 or 4(-5); rachilla with scattered short hairs; glumes 2.5-5 mm long, 2/5-1/2 the length of the spikelet, membranous to coriaceous, finely ciliate on margins, whitish-green, apex obtuse; lower glumes 2.5-3.5 mm long, oblong, 1-nerved; upper glumes 3.5-5 mm long, ovate, 3-nerved; lemmas 6-6.5(-7) mm long, lanceolate, 5-nerved, membranous, glabrous, whitish-green, sometimes purplish, awned, apex acuminate, the awn 1-2 mm long; callus glabrous; paleas 5/6 as long as the lemma, glabrous, ciliate on keels; lodicules ca. 0.8 mm long, lanceolate, acuminate; anthers 0.7-0.8 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 1/2-3/5 as long as the grain. Leaf blade anatomy.•Cross-sections with 5(-7) vascular bundles and 3(-5) ribs above; sclerenchyma present abaxially, continuous or discontinuous; bulliform cells absent; adaxial epidermis with few hairs, the hairs ca. 0.09 mm long. Observations.•Morphologically it appears that F boyacensis from the Colombian Cordillera Oriental is most similar to F chimborazensis. Other species that appear similar morphologically are F glumosa, F. imbaburensis, F. carchiense, and F sumapana. All of these species are more robust, i.e., with larger culms and have straight and often scabrous leaf blades. Distribution and habitat.•This species is endemic to Ecuador (Bolivar, Chimborazo, Cotopaxi, Pichincha, Tungurahua). It is known from humid or swampy patches in the superparamo growing between 3500^1500 m. 92 Festuca in South American Paramos 1 mm V. ! : .1 2 mm 2 cm 2 mm Figure 43. Festuca chimborazensis subsp. chimborazensis. A. Habit. B. Ligule. C. Inflorescence (detail). D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 3883 (PRC). Festuca in South American Paramos 0 -80 -75 200 93 400 Kilometers ( \ I -70 -65 A ; Figure 44. Distribution of Festuca chimborazensis (•), F. carchiense (A), F cleefiana (•), and F. dinirica (•). KEY TO THE SUBSPECIES OF FESTUCA CHIMBORAZENSIS 1. Leaf blades 4-9 cm long, curved; leaf cross-sections with only 5 vascular bundles; culms 8-15 cm tall; panicles 2-8 cm long, ca. 5 mm wide; upper glumes 4.5-5 mm long 34a. F. chimborazensis subsp. chimborazensis 1. Leafblades 11-15 cm long, mostly straight; leaf cross-sections with (5-)7 vascular bundles; culms 22-40 cm tall; panicles 8-10 cm long, ca. 7 mm wide; upper glumes 3.5-4 mm long 34b. F. chimborazensis subsp. micacochensis 34a. Festuca chimborazensis subsp. chimborazensis (Fig. 43). Culms 8-15 cm tall. Leafblades 4-9 cm long, curved; leaf cross-sections with only 5 vascular bundles. Panicles 2-8 cm long, ca. 5 mm wide. Spikelets with upper glumes 4.5-5 mm long. Additional specimens examined. ECUADOR. Bolivar: Road Ambato-Guaranda, 12 km W of intersection between old and new road, 01°26'S, 78°56'W, 4150 m, 1 Aug 1985, S. Laegaard 54820 (AAU, QCNE); 55.4 km SW of Ambato, 4050 m, 3 May 1990, P.M. Peterson 8979 & C.R. Annable (MO, QCA, QCNE, US); 66.5 km SW of Ambato, 94 Festuca in South American Paramos 4300 m, P.M. Peterson & C.R. Annable 8995 (MO, QCA, US); km 4 on the road Los Arenales-Salinas, 01°24'S, 78°55'W, S. Laegaard 55346 (AAU); 66.5 km SW of Ambato on Hwy to Guaranda and 2.2 km on road to Fecundo Vela, sandy volcanic soil in Festuca-Calamagrostis grassland, 4300 m, 5 Mar 1990, P.M. Peterson & C.R. Annable 8996A (MO). Chimborazo: Southern slope of Mt. Chimborazo, 3900 m, E. Asplund 8411, 8446 (NY, QCA, S); 3800 m, Fagerlind et al. 934bis (S); along Whymper road ca. 15 km S of Cruce de Los Arenales, 01°32'S, 78°52'W, 3950 m, 18 Sep 1998, S. Laegaard 19155 (AAU, QCNE); Urbina towards Mt. Chimborazo, 3700 m, E. Asplund 7892 (K, NY); sector Cruce de los Arenales, swampy, 01°28'14"S, 78°54'06"W, 4300 m, 20 Sep 2000, D. Stancik 3711, 3719 (PRC, QCA); 0P27'0.8"S, 78°53'57.5'W, 4150 m, 20 Sep 2000, D. Stancik 3710, 3720 (AAU, PRC, QCA). Cotopaxi: Volcan Cotopaxi, N side of the mountain, superparamo vegetation, sandy slope, 00°39'S, 78°42'W, 42504450 m, Sklendr & Kosteckovd 858 (AAU); Parque Nacional Cotopaxi, NW of Limpio Pungo, 00°37'S, 78°27'W, 3850-4000 m, S. Laegaard 52125 (AAU); Hacienda Pauzacha S of Volcan Cotopaxi, 00°44'S, 78°49'W, 3650 m, S. Laegaard 55736 (AAU); along road El Chaupi-Pastocalle, 00°41 'S, 78°39'W, 3500 m, 2 Jul 1985, S. Laegaard 54594 (AAU, QCA, QCNE); Angamarca road 5 km from junction to road Latacunga-La Mana, 01°00'S, 78°55'W, 4250-4280 m, S. Laegaard 102156 (AAU); 21.5 km E of Pilalo, 3820 m, 15 Apr 1990, P.M. Peterson 8768, C.R. Annable & M.E. Poston (MO, QCA, US); Tililac, 4000 m, Acosta-Solis 16765 (US); Mun. Chaupi, NE slope of Volcan Illiniza Norte, 00°38'S, 78°42'W, 4000^1050 m, 12 Oct 2000, D. Stancik 4024, 403IB, 4033B (PRC, QCA); swampy patches, 00°38'S, 78°42'W, 4000-4050 m, 12 Oct 2000, D. Stancik 4034 (AAU, PRC, QCA, US); Mun. Lasso, Parque Nacional Cotopaxi, NNE slope of Volcan Cotopaxi, valley of Quebrada Taniloma, 00°36.8'S, 78°24.3'W, 3750 m, 21 Jun 1999, D. Stancik3080 (AAU, PRC, QCA, W); E slope of Volcan Cotopaxi, 00°38'S, 78°25'W, 4300 m, 28 Sep 2000, D. Stancik 3880 (AAU, PRC, QCA); plain below SE slope of Volcan Cotopaxi, 0°37'S, 78°24'W, 3800-3900 m, 28 Sep 2000, D. Stancik 3883 (PRC, QCA). Pichincha: Iter Regnellianum quartum, Flora Aequatoriensis, prov. Pichincha, Sincholagua, 4200 m, 7 Jul 1939, E. Asplund 8641 (S-paratype); mountain ridge ca. 2 km to the W from Cerro Saraurcu, 4200-4350 m, Sklendr & Kosteckovd 1322 (AAU); Volcan Iliniza, NE slope below the refugio, 00°39'S, 78°42'W, 4430 m, Holm-Nielsen 24871 (AAU); Entre Pifo y boqueron de Cerro Corrales, Paramo de Guamani, 4050 m, Barclay & Juajibioy 8905 (COL, MO); Volcan Antisana, between campsite IMAP and Laguna Micacocha, 00°33 'S, 78°12'W, 3850-3950 m, S. Laegaard 101583 (AAU, QCA, QCNE); N side, ca. 12 km along road from Hacienda Antisana, 00°27'S, 78°10'W, S. Laegaard 102892 (AAU); 00°30'S, 78°00'W, 4100 m, Grubb et al. 620 (NY, US); NE slope of Rucu Pichincha, 00°10'S, 78°34'W, 4300 m, Sklendr & Kosteckovd 1-24 (NY). Tungurahua: Comunidad Rumipata, pasture in paramo, 00°22'S, 78°55'W, 4000 m, Brandbyge 42597 (AAU, MO). 34b. Festuca chimborazensis subsp. micacochensis Stancik, Folia Geobot. Phytotax. 39(1): 105, f. 3, 6-10. 2004. TYPE: Ecuador. Pichincha/ Napo, Volcan Antisana, between Campamento IMAP and Laguna Micacocha, 00°33'S, 78°12'W, naked soil on trail, 3850-3950 m, 7 Mar 1992, S. Laegaard 101612 (holotype: AAU!; isotypes: PRC!, QCA!, QCNE!). Culms 22-AQ cm tall. Leaf blades 11-15 cm long, mostly straight; leaf cross-sections with (5-)7 vascular bundles. Panicles 8-10 cm long, ca. 7 mm wide. Spikelets with upper glumes 3.5^4 mm long. Distribution and habitat.•Festuca chimborazensis subsp. micacochensis has so far only been recorded from three sites: western slopes of Antisana, northern slopes of Chimborazo, and at Guagua Pichincha. At all three sites this subspecies is sympatric with F chimborazensis subsp. chimborazensis. Additional specimens examined. ECUADOR. Bolivar: 55.4 km SW of Ambato on Highway to Guaranda, 4050 m, 3 May 1990, P.M. Peterson 8982 & C.R. Annable (K, MO, QCA, QCNE, US); km 42.7,4020 m, 3 May 1990, P.M. Peterson 8973 & C.R. Annable (MO, QCA, QCNE, US); along main road ca. 1 km W of Cruce de Los Arenales, 01°25'S, 78°54'W, 4050-4100 m, S. Laegaard 101517 (AAU). Pichincha: Pifo-Pintag road, in valley 2.5 hours horseride above Inga Monserrat, 00°19'S, 78°17'W, 3625-3725m, 11 Apr 1992, S Laegaard 102253 (AAU, QCA, QCNE); road Lloa-Guagua Pichincha, km 11,00°12'S, 78°35'W, 4300 m, S. Laegaard et al. 102744 (AAU); S. Laegaard et al. 102739 (AAU, QCA). Festuca in South American Paramos 35. Festuca chita Stancik, Darwiniana 41(1^4): 129, f. 13a-f. 2003. (Figs. 45, 49). TYPE: Colombia. Boyaca, Canon del Chicamocha, Mun. Chita, Vereda Los Colorados, 3300 m, 14 May 1991, Etter 661 (holotype: COL!). Tussocked perennials with intravaginal innovations. Culms 40-60 cm tall, erect, glabrous; nodes 1, basal; cataphylls small, coriaceous. Leaf sheaths membranous, brown to purplish-brown, glabrous; auricles absent; ligules 0.3-0.5 mm long, membranous, apex truncate, short-ciliate; blades 10-15 cm long, 0.4-0.6 mm wide, involute, green, abaxially glabrous, apex obtuse. Panicles ca. 8 cm long, ca. 3 cm wide, contracted; branches glabrous. Spikelets 8-9 mm long, lanceolate, florets 3 or 4; rachilla densely hairy; glumes 5.5•8(•8.5) mm long, coriaceous, dark purple; lower glumes 5.5-6.5 mm long, narrowly lanceolate, 1-nerved; upper glumes 7•8(•8.5) mm long, lanceolate, 3-nerved, sparsely scabrous; lemmas 6.5-7.5 mm long, lanceolate, 5-nerved, coriaceous, dark purple, scabrous and short-hairy, entire, awned, the awn 0.5-1.5 mm long; callus glabrous or sparsely hairy; paleas 2/3-4/5 as long as the lemma, glabrous, apex hairy; anthers 0.8-1.1 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 1/2 as long as the grain. Leaf blade anatomy.•Cross-sections with 7-11 vascular bundles and 5 ribs, sclerenchyma under abaxial epidermis discontinuous, extending to all the vascular bundles, adaxial sclerenchyma present, extending to vascular bundles only exceptionally; bulliform cells absent; adaxial epidermis hairy, the hairs ca. 0.5 mm long. Geographical distribution.•This species is endemic to the Colombian Cordillera Oriental, at Canon del Chicamocha and is only known from the type locality at an altitude of 3300 m. The Canon del Chicamocha is a dry valley without typical paramo vegetation. Additional specimens examined. COLOMBIA. Boyaca: Canon del Chicamocha, Mun. Chita, Vereda Los Colorados, 3300 m, 27 Jan 1927, Killip 18479 (COL). 36. Festuca cleeflana E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 69: 1548.1984. (Figs. 44, 46, 90C-F). TYPE: Colombia. Boyaca, Paramo de la Rusia, NW-N de Duitama Aislada, 3580 m, 7 Dec 1972, A.M. Cleef 6826 (holotype: US-2785718!; isotypes: COL!, U!). 95 Densesly tussocked perennials with intravaginal innovations. Culms 70•130(•150) cm tall, erect, glabrous sometimes finely scabrous; nodes 1 or 2(-3). Leaf sheaths, membranous to coriaceous, grayish-brown, glabrous; auricles absent; ligules (0.5-)l-2(-2.5) mm long, membranous to coriaceous, apex mostly obtuse, short-ciliate; blades 30-50(-60) cm long, 0.6-0.9(-l. 1) mm wide, conduplicate, abaxially glabrous, green, apex obtuse. Panicles 10-20(-25) x 3-10(-15) cm, flexuous, branched, erect, oblong; branches finely scabrous. Spikelets 8-11 mm long, oblong, florets 4-6(-7); rachilla long, densely pilose; glumes (3.5-)4-7.5 (-8) mm long, narrowly lanceolate, coriaceous, purple or purplish-green, upper 1/4 scabrous, apex acute; lower glumes (3.5-)4-6(-6.5) mm long, 1-nerved; upper glumes (5.5-)6•7.5(-8) mm long, 3-nerved; lemmas 6-7.5(-8) mm long, lanceolate, membranous to coriaceous, 5-nerved, purplishgreen, scabrous to densely hairy, apex short twodentate, awned, the awn 0.5-2.5 mm long; paleas as long as the lemma, papillose, upper 1/3 hairy; lodicules oblong-lanceolate, acuminate; anthers (2-)2.5-3 mm long; ovary apex glabrous. Caryopses oblong-lanceolate; hilum 3/4 as long as the grain, linear. Leaf blade anatomy.•Cross-sections with 7•9(•11) vascular bundles and 5-9 ribs; sclerenchyma under abaxial epidermis continuous or discontinuous, extending to all the vascular bundles; adaxial epidermis present, extending to all or every other vascular bundle; bulliform cells absent; adaxial epidermis densely hairy, the hairs 0.02-0.07 mm long. Observations.•Festuca cleefiana is an easily recognizable species, forming relatively large tufts with flexuous, ramified panicles and spikelets with typically long glumes. Distribution and habitat.• Festuca cleefiana is endemic to Colombian Cordillera Oriental (Cundinamarca, Boyaca, Santander). It is a typical species of the grass paramo occurring between 3300-4000 m. This species is known from the communities of Festuca dolichophylla & Paspalum bonplandianum (Rangel & Sturm 1995), Sphagno magellanici-Chusquetum tesellatae (Rangel & Sturm 1995), Hypericum juniperinum (Cleef 1981). Additional specimens examined. COLOMBIA. Cundinamarca: Mun. Bogota, valley between Usme and Nazareth, 4°13'N, 74°H'W, 3480 m, 16 Jul 1998, D. Stancik272 (PRC); Mun. 96 Festuca in South American Paramos Figure 45. Festuca chita. A. Stylized growth form. B. Habit. C. Spikelet. D. Glumes. E. Lemma. F. Leaf blade cross-section. A-F, Etter 661 (COL). Festuca in South American Paramos 97 2 mm Figure 46. Festuca cleefiana. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 1421 (PRC). Villapinzon, Paramo La Calera, 5°12'N, 73°33 'W, 3500 m, 22 Jul 1978, D. Stancik 409, 466, 467 (COL, PRC). Boyaca: Mun. Aquitania, Paramo El Guarne, 3500 m, 8 Feb 1999, D. Stancik 2160, 2188, 2189 (COL, FMB, COL); Mun. Aquitania, road to Sisvaca, km 6, 2900 m, 10 Feb 1999, D. Stancik 1920 (COL, FMB, PRC); D. Stancik 2257 (COL, FMB, PRC); km 2, 3350 m, 14 Jan 1999, D. Stancik 1950 (PRC); 3240 m, 14 Jan 1999, D. Stancik 1964 (COL, FMB, PRC); 3450 m, 14 Jan 1999, D. Stancik 1935 (COL, FMB, PRC); Mun. Aquitania, Paramo Sarna, 3400 m, Festuca in South American Paramos 20 Jan 1999, D. Stancik 2000 (COL, PRC); Mun. Arcabuco, Santuario Iguaque, Laguna Iguaque, 3600 m, 14 Oct 1998, D. Stancik 924, 926, 945 (COL, FMB, PRC); Lagun Ojo de Agua, 3650 m, 14 Oct 1998, D. Stancik 927 (COL, FMB, PRC); Mun. Arcabuco, SW slope of Cerro Volcan, 3000 m, 20 Nov 1998, D. Stancik 1407 (COL, FMB, PRC); Mun. Belen, Paramo de Guina, 3250 m, 6 Feb 1999, D. Stancik 2130, 2136 (COL, PRC); 3300 m, 6 Feb 1999, D. Stancik2137, 2138 (PRC); Mun. Duitama, Paramo de La Rusia, 5°54'27"N, 73°4'44"W, D. Stancik 1004 (COL); 3750 m, 12 Feb 1999, D. Stancik 2367, 2372, 2373 (COL, PRC); 3580 m, 3540 m, Hernandez 1311 (COL); Paramo Pan de Azucar, 3400 m, 30 Nov 1998, D. Stancik 1421, 1422 (COL, FMB, PRC); 3350 m, 30 Nov 1998, D. Stancik 1423 (COL, FMB, PRC); 3400 m, 30 Nov 1998, D. Stancik 1482 (COL, FMB, PRC, US); Mun. El Cocuy, Parque Nacional El Cocuy, Las Cabanas Kanwara, 4070 m, 30 Dec 1998, D. Stancik 1827, 1865 (COL, FMB, PRC); 4350 m, 30 Dec 1998, D. Stancik 1828, 1830 (COL, FMB, PRC); pass between Cocuy and Chita, 4000 m, 3 Feb 1985, Wood 5141 (COL, K); Mun. Chinavita, Cerro Mamapacha, 2900 m, 11 Dec 1998, D. Stancik 1602, 1635, 1640 (COL, FMB, PRC); 2850 m, 11 Dec 1998, D. Stancik 1629 (COL, PRC); 11 Dec 1998, D. Stancik 1576, 1577, 1628 (COL, FMB, PRC); 11 Dec 1998, D. Stancik 1580 (COL, PRC); Mun. Mongui, Laguna La Colorada, 3550 m, 21 Jan 1999, D. Stancik 2020 (COL, FMB, PRC); 3650 m, 21 Jan 1999, D. Stancik 2045 (COL, FMB, PRC); Mun. Paipa, Cuchilla El Paramo, 3200 m, 4 Dec 1998, D. Stancik 1503, 1532, 1555, 1556 (PRC); Mun. Santa Rosa de Viterbo, Alto Lamadero, 3200 m, 30 Nov 1998, D. Stancik 1438, 1457, 1463, 1465 (COL, PRC); Mun. Siachoque, Paramo Siachoque, 05°30'N, 73°8'W, 3650 m, 24 Jan 1999, D. Stancik 2051, 2050, 2054 (COL, FMB, PRC); Mun. Socota, Paramo Pisva, lagoon Chorro Negro, 3500 m, 11 Feb 1999, D. Stancik 2346, 2350 (COL, FMB, PRC); Alto de Cardon, 3540 m, 11 Feb 1999, D. Stancik2314 (COL, FMB, PRC); Alto de Calarca, 3500 m, 11 Feb 1999, D. Stancik 2313, 2332 (COL, FMB, PRC); Mun. Sotaquira, Paramo Chontales, 3050 m, 15 Dec 1998, D. Stancik 1688, 1689, 1690 (COL, FMB, PRC); 14 Nov 1998, D. Stancik 1339 (COL, FMB, PRC); Mun. Pesca Paramo Cortadero, vereda Puerta Chiquita, 3500-3700 m, 15 Oct 1981, Bejarano 44 (COL); Paramo de Belen, 3500 m, 15 Nov 1999, Rangel etal. 3515 (COL); Mun. Toca, Paramo Cortadero, 5°30'N,73°15'W, 3450m, 14Nov 1998,/). Stancik 1339 (COL, FMB, PRC); 3350 m, 14 Nov 1998, D. Stancik 1366 (COL, FMB, PRC); 3600 m, 14 Nov 1998, D. Stancik 1373 (COL, FMB, PRC); Mun. Labranzagrande, Paramo Franco, 3300 m, 9 Feb 1999, D. Stancik 2224, 2230, 2232, 2236, 2240 (COL, FMB, PRC); Mun. Labranzagrande, Valle de Rio Cushiana, 5 km from Toquilla, 2950 m, 9 Feb 1999, D. Stancik 2249 (COL, FMB, PRC). Santander: Mun. Concepcion, vereda Juradito, Paramo de Gallina, 3500 m, 24 Feb 1999, D. Stancik 2495, 2502, 2503, 2505, (COL, FMB, PRC); 24 Feb 1999, D. Stancik 24S9 (PRC). 37. Festuca cocuyana Stancik, Darwiniana 41 (1-4): 133, f. 12a-e. 2003. (Figs. 39, 47, 91A-D). TYPE: Colombia. Boyaca, Mun. Cocuy & Guican, Parque Nacional El Cocuy, Alto de las Cuevas. Grass paramo with Calamagrostis effusa, Espeletia sp., Acaulimlva sp., etc., 3850 m, 30 Dec 1999, D. Stancik 1886 (holotype: PRC!; isotypes: COL!, FMB!). Tussocked perennials with intravaginal innovations. Culms 20-50 cm tall, erect, glabrous; nodes 1, basal. Leaf sheaths membranous to coriaceous, stramineous, glabrous, finely striate; auricles absent; ligules 0.6-1 mm long, membranous to coriaceous, apex truncate; blades 15-20 cm long, 0.8-1.5 (-2) mm wide, conduplicate or flat, abaxially glabrous. Panicles 8-12 cm long, 0.8-1.5 mm wide, contracted, narrow, elongate. Spikelets 10-13 mm long, lanceolate, florets 2 or 3; rachilla pubescent; glumes 7.5-10.5 mm long, almost as long as the spikelet, lanceolate, membranous, scabrous along midnerve, apex acute; lower glumes 7.5-8 mm long, 1-nerved; upper glumes 7.5-10.5 mm long, 3-nerved; lemmas 8-10 mm long, lanceolate, 5-nerved, membranous, awnless or short-awned, the awn ca. 0.5 mm long; callus glabrous or sparsely hairy; paleas 3/4 as long as the lemma, papillose, distally scabrous; lodicules 1-1.4 mm long, lanceolate, two-dentate; anthers 0.8-1.1 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 3/4 as long as the grain, linear. Leaf blade anatomy.•Cross-sections with (8-) 10-11 vascular bundles and 5-7 ribs above; sclerenchyma under both abaxial and adaxial epidermis, discontinuous; sclerenchyma girders absent; bulliform cells absent; adaxial epidermis with scattered hairs, the hairs ca. 0.02 mm long. Festuca in South American Paramos 99 1 mm Figure 47. Festuca cocuyana. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 1886 (PRC). 100 Festuca in South American Paramos Observations.•Festuca cocuyana is morphologically similar to F. sanctae-martae, an endemic species of Sierra Nevada de Santa Marta. Festuca cocuyana differs from F sanctae-martae by having shorter culms (20-50 verses 60-80 cm), shorter leaf blades (15-20 verses 25-35 cm), and shorter lower glumes (7.5-8 verses 8.5-9.5 mm). Distribution and habitat.• It is endemic to the Sierra Nevada del Cocuy in the Colombian Cordillera Oriental. It occurs between 3800^1300 m, in grass paramo dominated by Calamagrostis effusa and Espeletia spp., and on rocky slopes in the superparamo. Additional specimens examined. COLOMBIA. Arauca: Sierra Nevada del Cocuy, cabeceras de la Quebrada El Playon, Patio Bolos 2.5 km, al NW de la Laguna La Plaza, hoya S. Jose, 4300 m, 10 Mar 1973, Cleef9078 (COL); S of the Lagoon La Plaza, 4300 m, 31 Dec 1985, Wood 5260 (COL, FMB, K). 38. Festuca cundinamarcae E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 69: 1548. 1984. (Figs. 48,49,91E & F, 92A& B). TYPE: Colombia. Cundinamarca, Macizo de Bogota, Cerro Diego Largo, paramo, 3540 m, 14 Jan 1940, J. Cuatrecasas 7966 (holotype: S!; isotypes:COL!,U!,US!). Tussocked perennials with intravaginal innovations. Culms 60-100(-120) cm tall, erect, scabrous; nodes 1 on distal half. Leaf sheaths coriaceous, stramineous, glabrous; ligules 0.1-0.3 mm long, coriaceous, apex truncate, short-ciliate; blades 30-50 cm long, 0.7-1 (-1.2) mm wide, conduplicate, rigid, abaxially glabrous, dark green, apex obtuse ormucronate. Panicles 10-20 x 0.5-1.5 cm, contracted, narrow; branches scabrous. Spikelets 9-13 mm long, oblong-lanceolate, florets 5•7(•8); rachilla sparsely short-pilose; glumes 36 mm long, lanceolate, membranous to coriaceous, green, scabrous distally, apex acute; lower glumes 3^4.5 mm long, 1-nerved; upper glumes (4.5-)5-6 mm long, 3-nerved; lemmas (5.5-)6-7(-7.5) mm long, lanceolate, 5-nerved, membranous to coriaceous, glabrous, upper 1/3 scabrous, apex entire, usually awned, the awn up to 1 mm long; paleas as long as the lemma, papillose, upper 1/3 hairy; lodicules ca. 1.1 mm long, lanceolate, acuminate; anthers 2-3 mm long; ovary apex glabrous. Caryopses oblong-lanceolate; hilum 2/3 as long as the grain, linear. Leaf blade anatomy.•Cross-sections with 9-15 vascular bundles and 7-13 ribs; sclerenchyma under abaxial epidermis discontinuous, extending to all or every other the vascular bundles, adaxial sclerenchyma present, often extending to the vascular bundles forming girders; bulliform cells absent; adaxial epidermis densely hairy, the hairs 0.09-0.2 mm long. Distribution and habitat.•Festuca cundinamarcae is endemic to Colombia, where it is known only from the Cordillera Oriental (Cundinamarca, Boyaca, Meta). It is a common species of grass paramos growing between 2700-3700 m. This species occurs in communities such as: Pentacalio nitidae-tessellatae (Sanchez & Rangel 1990), Calamagrostietum planifoli-effusae (Vargas & Zuloaga 1985), Espeletietum argenteaeCalamagrostiosum effusum (Cuatrecasas 1934), Bartsio santolinifoliae-Calamagrostietum effusae (Rangel & Ariza 2000). Additional specimens examined. COLOMBIA. Boyaca: Boyaca, 2700 m, 15 Apr 1964, Saravia 3912 (COL, PRC); Mun. Pesca, Puerta Chiquita, Paramo de Cortadera, 3500-3700 m, 15 Oct 1981, Bejarano 22 (UPTC); Mun. Aquitania, road SuseSisvaca, km 1.5-2,3050 m, 13 Jan 1999,/). Stancik 1939,1951,1952 (COL, FMB, PRC); 14 Jan 1999, D. Stancik 1944 (PRC); Paramo Sarna, 3350 m, 20 Jan 1999, D. Stancik 1987, 1988 (COL, FMB, PRC); 3400 m, 20 Jan 1999, D. Stancik2005 (COL, FMB, PRC); 3200 m, 20 Jan 1999, D. Stancik 1985, 1986 (COL, FMB, PRC); Mun. Aquitania, Paramo Suse, 3350 m, 14 Jan 1999, D. Stancik 1929 (COL, PRC); 14 Jan 1999, D. Stancik 1945 (PRC); 14 Jan 1999, D. Stancik 1922 (COL, FMB, PRC); Mun. Belen, Paramo Giiina, 3250 m, 6 Feb 1999, D. Stancik 2109, 2115, 2116, 2117, 2120 (COL, PRC); 6 Feb 1999, D. Stancik 2114, 2136 (COL, FMB, PRC); Mun. Mongui, Laguna La Colorada, 3300 m, 21 Jan 1999, D. Stancik2019 (COL, FMB, PRC); Mun. Toca, Paramo Cortadero, 05°30'N, 73°15'W, 2700 m, 14 Nov 1998, D. Stancik 1402 (COL, FMB, PRC); 3200 m, 14 Nov 1998, D. Stancik 1347,1360,1363, /596(COL,FMB,PRC); 4 Dec 1998, D. Stancik 1513 (COL, FMB, PRC); Mun. Samaca, Paramo Rabonal, 2000 m, 1 Nov 1998, D. Stancik 1286 (COL, FMB, PRC); Mun. Toquilla, Valle of Rio Cushiana, 2950 m, 9 Feb 1999, D. Stancik 2250 (COL, FMB, PRC); Mun. Tutasa, Paramo de Carnicerias, 3300 m, 26 Feb 1999, D. Stancik 2597, 2599, 2612 (PRC); Mun. Paipa, cuchilla El Paramo, 3250 m, 3 Dec 1998, Festuca in South American Paramos 101 Figure 48. Festuca cundinamarcae. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 3539 (PRC). 102 Festuca in South American Paramos -80 -75 -70 -65 Figure 49. Distribution of Festuca glumosa (•), F. cundinamarcae (A), F. chita (•), and F.turimiquirensis (•). D. Stancik 1523 (COL, FMB, PRC); 4 Dec 1998, D. Stancik 1511 (COL, PRC); Mun. Susacon, km 20, margin of river Susacon, 3150 m, 23 Feb 1999, D. Stancik 2472 (PRC); 23 Feb 1999, D. Stancik 2471, 2473 (COL, FMB, PRC). Cundinamarca: Cordillera de Bogota, 2900 m, Nov 1856, Triana s.n. (US); Bogota, Paramo de Monserrate, vereda El Verjon, 3000-3200 m, 29 Jul 1988, Garzon 617 (COL); 3160 m, 15 Feb 1980, Zuluoga 107 (COL); 3000-3300 m, 16 Apr 1944, Killip et al. 38050 (COL); 3125m, 24 May 1968, Valderrano 34 (UPTC); 3125m, 26 Apr 1968, Sturm 33 (UPTC); Mun. Bogota, Alto de la Viga, 1 Feb 2000, D. Stancik 3539 (COL, PRC); Paramo de Cruz Verde, 3400-3500 m, 15 Sep 1940, J. Cuatrecasas 10446 (COL, US); 04°34'N, 74°01'W, 3500-3700 m, 17 Jun 1995, S. Laegaard & Mayorga 17536 (AAU, COL); Sumapaz-Santa Rosa, 3600 m, 15 Jul 1984, Wood 4539 (COL, FMB); Hoya de Quebrada Sitiales, 1.5 km NW de Laguna La Guitarra, 26 Jan 1972, Cleef 1090 (COL, U); Paramo Chingaza, sector Rio La Playa, Valle de Fraillejon, 3250 m, 14 Nov 1992, Guerrero 152 (COL); near La Calera, 3500 m, 1983, Wood 3523 (AAU, COL, K); Paramo de Palacio, La Siberia, 3400 m, 30 Feb 1961, Rodriguez 4130 (COL); Paramo between Bogota and Choachi, 3320 m, 7 Jan 1974, G. Davidse et al. 5545 (COL, US); Paramo entre Cogua y San Cayetano, cercania de la Laguna Seca, 3700 m, 12 Feb 1972, Cleef 6285 (COL); Paramo de Guasca, 3200-3300 m, 2 Jun 1940, J. Cuatrecasas 9475 (COL, US). Meta: Paramo Sumapaz, Hoya Sitiales, laguna La Primavera, 3580 m, 25 Jan 1972, Cleef 982A (COL). 39. Festuca densipaniculata E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 69: 1551. 1984. (Figs. 50, 52, 92C-F). TYPE: Ecuador. Festuca in South American Paramos 103 2 mm Figure 50. Festuca densipaniculata. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Asplund 8397 (S). 104 Festuca in South American Paramos Chimborazo, southern slope of Mt. Chimborazo, sandy ground, 4600 m, 18 Mar 1939, E. Asplund 8397 (holotype: S!). Tussocked perennials with intravaginal innovations. Culms 20-50 cm tall, erect, finely scabrous; nodes 1, basal. Leaf sheaths coriaceous, stramineous, glabrous or scabrous; ligules 0.8•1.5(•1.7) mm long, membranous, apex truncate, short-ciliate; blades 17-25 cm, long, 0.8-1.2 mm wide, conduplicate, abaxially scabrous, glaucous, apex pointed. Panicles 5-10 x 1.5-2 cm, contracted, dense, lanceolate to oblong; branches finely scabrous throughout. Spikelets 8•9(•10) mm long, elliptic to oblong, florets 3 or 4; rachilla densely hairy; glumes 4•6 mm long, membranous to coriaceous, green and somewhat glaucous, apex acute; lower glumes 4-5 mm long, lanceolate to oblong-lanceolate, 3-nerved, glabrous or upper 1/3 scabrous; upper glumes (4.5-)5-6 mm long, oblong to oblong-lanceolate, 5-nerved, glabrous or upper 1/2 scabrous; lemmas (6.5-)7-8.5 mm long, lanceolate, 5-7- nerved, membranous, green or sometimes purplish, glaucous, upper 1/2 or entire surface scabrous, awned, the awn 0.7-1 mm long; callus glabrous; paleas as long as the lemma, lanceolate, membranous, upper 1/3 and along keels scabrous to hairy; lodicules 1.4•2 mm long, oblong; anthers (2.3-)2.7-2.8 mm long; ovary apex glabrous. Caryopses lanceolate to oblong-lanceolate; hilum 1/2 as long as the grain, linear. Leaf blade anatomy.•Cross-sections with (7-) 11-15 vascular bundles and 5-7 ribs above; sclerenchyma under abaxial epidermis continuous, adaxial sclerenchyma present, extending to every other vascular bundle forming girders; abaxial epidermis with a dense covering of prickles, adaxial epidermis with frequent hairs, the hairs 0.2-0.4 mm long. Observations.•The type specimen of F. densipaniculata was originally annotated by Alexeev as 'Festuca densiflora" but he published it as F. densipaniculata because of the earlier homonym, F. densiflora Tovar. There are only two known collections of this species. Leaf anatomical characteristics suggest that F subulifolia and F asplundii may be closely related. Otherwise, the status of this species is not readily apparent and molecular studies will be needed for elucidation of its affinities. Distribution and habitat.•Festuca densipaniculata is known only from grass paramos of central Ecuador between 4000^1600 m. Additional specimens examined. ECUADOR. Chimborazo: S slope of Mt. Chimborazo, sandy ground, 4600 m, E. Asplund 8396 (S-paratype). Napo: Antisana, 22 Jul 1997, Sklendf & Kosteckova 2738 (PRC). 40. Festuca dinirica Stancik, Novon 14(3): 341-343, f. If•j. 2004. (Figs. 44, 51). TYPE: Venezuela. Lara, Mun. Humocaro Alto, Parque Nacional Dinira, 09°35'39"N, 70°07'12"W, grass paramo with shrubby patches with Rhynchospora sp., Chusquea sp., Blechnum sp., Calamagrostis effusa, Espeletia sp., and Diplostephium sp., 3170 m, 30 Nov 2000, D. Stancik 4287 (holotype: PRC!; isotypes: CAR!, COL!). Tussocked perennials with intra- and extravaginal innovations. Culms 50-60 cm tall, erect, glabrous; nodes 1, in distal half. Leaf sheaths coriaceous, brown, glabrous, fibrous near base; auricles absent; ligules 0.5-1 mm long, membranous, apex truncate; blades 20-25 cm long, 0.4-0.6 mm in diameter, conduplicate to involute, abaxially glabrous, green. Panicles 10-12 x 0.5-1.5 cm, contracted, erect; branches glabrous. Spikelets 8-10 mm long, lanceolate sometime oblonglanceolate, florets 3 or 4; rachillas 1-1.2 mm long, sparsely hairy; glumes 3.4•6 mm long, lanceolate, membranous to coriaceous, violet, apex acute; lower glumes 3.4-4 mm long, 1-nerved; upper glumes 5.5-6 mm long, 3-nerved; lemmas 6-7 mm long, lanceolate, 5-nerved, coriaceous, violet, scabrous, awned, the awn 1.5•4 mm long; callus glabrous; paleas as long as the lemma, lanceolate, membranous, scabrous; anthers 2-2.5 mm long; ovary apex glabrouse. Caryopses lanceolate; hilum 4/5 as long as the grain, linear. Leaf blade anatomy.•Cross-sections usually with 5 vascular bundles and 3-5 ribs above; sclerenchyma under abaxial and adaxial epidermis discontinuous; adaxial epidermis hairy, the hairs ca. 0.09 mm long. Observations.•The spikelet structure of F dinirica resembles F. tolucensis, another frequent species of the Venezuelan paramos. However F dinirica clearly differs from F tolucensis by its smaller culms (40-60 versus 60-80 cm), mixed innovations (versus intravaginal), brown, striate, and fibrous leaf sheaths (versus stramineous and non-fibrous), shorter ligules (0.5-1 mm versus 1.8-3.5 mm), and anatomically with discontinuous Festuca in South American Paramos 105 2 mm Figure 51. Festuca dinirica. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf blade cross-section. A-G, Stancik 4288 (PRC). Festuca in South American Paramos 106 -10 -10 200 -80 -75 -70 400 600 Kilometers A -65 Figure 52. Distribution of Festuca glyceriantha (•), F. holubii(A), F. densipaniculata (•), and F. monguensis (•). abaxial sclerenchyma and all vascular bundles free (versus continuous abaxial sclerenchyma with some girders persent). The characteristics of mixed innovations with brown, striate, fibrous sheaths in F. dinirica suggest a close relationship with F. sect. Aulaxyper. In South America, this section is not very common and is still poorly studied. The Colombian and Ecuadorian species, F. andicola, is geographically the nearest species within F. sect. Aulaxyper. Distribution and habitat.•Festuca dinirica is endemic to Venezuela (Lara) and occurs at an altitude of about 3200 m where it forms small tussocks in shrubby-grassy paramos and is associated with various species of Blechnum, Espeletia, Hypericum, Calamagrostis, Chusquea, and Rhynchospora. Additional specimens examined. VENEZUELA. Lara: Mun. Humocaro Alto, Parque National Dinira, 09°35'39"N, 70°07'12"W, 3170 m, 30 Nov 2000, D. Stancik 4288 (CAR, COL, PRC). 41. Festuca glumosa Hack, ex E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 69(11): 1549. 1984. (Figs. 49, 53, 93A-D). TYPE: Ecuador. In pasq. andinis, 1886, A. Sodiro 36/4 (holotype:W!) Festuca ovina subvar. jamesonii St.-Yves, Candollea 3: 166. 1927. TYPE: Ecuador Andes de Quito, 15000 ft, Jameson 230 (lectotype: G-DC!, designated here). Densely tussocked perennials with intravaginal innovations. Culms 15-55 cm tall, erect, finely scabrous; nodes 1, basal with 2 leaves. Leaf sheaths membranous to coriaceous, firm, stramineous glabrous, inconspicuously striate; ligules 0.8-1.2 mm long, apex two-dentate, short-ciliate; blades 10-25 cm long, 0.8-1.4 mm wide, involute, abaxially scabrous, glaucous, apex obtuse. Panicles 8-10 x 0.6-1 cm, linear to lanceolate, Festuca in South American Paramos 107 1 mm Figure 53. Festuca glumosa. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 3392 (PRC). 108 Festuca in South American Paramos contracted, dense, branches scabrous. Spikelets 8.5-10 mm long, lanceolate, florets 3 or 4; rachilla with scattered hairs; glumes 4.5-6.5 mm long, lanceolate, membranous to coriaceous, keeled, purplish-green, upper 1/3 scabrous along keel, apex acute; lower glumes 4.5-6 mm long, 1-nerved; upper glumes 6-6.5 mm long, 3-nerved; lemmas 7-8 mm long, lanceolate, 5-nerved, membranous, keeled, purplish-green, scabrous distally, awned, the awn 0.5-1.5 mm long; callus with scattered hairs; paleas 4/5 as long as the lemma, lanceolate, membranous, upper 1/3 and keels scabrous to hairy; lodicules lanceolate; anthers 0.8-0.9 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 1/2-3/5 as long as the grain, linear. Characters Leaf width (mm) Leaf length (cm) Ligule length (mm) Panicle length (cm) Spikelet length (mm) Number of florets Lower glume length (mm) Upper glume length (mm) Lemma length (mm) Awn length (mm) Anther length (mm) F. glumosa 0.8-1.4 10-25 0.8-1.2 8-10 8.5-10 3 or 4 4.5-6 6-6.5 7-8 0.5-1.5 0.8-0.9 Distribution and habitat.•Festuca glumosa ranges from central Ecuador to the northern Cordillera Central in Colombia and occurs in (dry) grasslands and rocky slopes of superparamos between 4000-4600 m. Festuca glumosa is known from the following communities: Diplostephio eriophori-Loricarietum colombianae (Salamanca et al. 1991), Cerastio floccosi-Pentacalietum gelidae (Salamanca et al. 1991), Lupino alopecuroidisAgrostietum araucanae (Salamanca et al. 1991), Baccharido caespitosae-Agrostietum araucanae (Salamanca et al. 1991), and Aciachne acicularisCalamagrostis intermedia (Verweij 1995). Additional specimens examined. COLOMBIA. Caldas: Mun. Manizales, Parque Nacional Los Nevados, between Crater La Oletta and el Ruiz, way to Rio Nereidas, 4200-4500 m, 18 Sep 1999, D. Stancik 3393 (COL, PRC, W); D. Stancik 3392 (COL, PRC); D. Stancik 3413 (COL, PRC); 19 Sep 1999, Casa del Cisne, 4180 m, 19 Sep 1999, D. Stancik 3383 (COL, PRC); way from Rio Nereidas, km 5-7,4000 m, 18 Leaf blade anatomy.•Cross-sections with 7 (rarely 5) vascular bundles and 5 ribs above; sclerenchyma usually discontinuous under abaxial epidermis and small or absent under adaxial epidermis; adaxial epidermis with scattered hairs, the hairs ca. 0.09 mm long. Observations.•A specimen of F. glumosa labeled as Sodiro 36/4 was found at QPLS. However, this specimen is not the type specimen since the collection date is Sep 1887 and the locality is "crescit in pasquis on Pichincha". Apparently, Sodiro used a single number to indicate the same species. Morphologically similar species are F. carchiense and F. imbaburensis. The species may be distinguished by the characters shown in the following tabular comparison. F. carchiense 0.4-0.7 25^10 0.5-0.8 10-20 9-10.5 3or4 4-4.5 5-6.5 7-7.5 0.8-1.5 1.1-1.3 F. imbaburensis 0.5-0.6 9-20 0.5-0.7 6-12 8-10 2 or 3 3-3.5 4-4.2 5.5-6 0.5-0.7 1-1.5 Sep 1999, D. Stancik 3394 (COL, PRC); Mun. Villamaria, Crater Olleta, 04°54'N, 75°21 'W, 4600 m, 27 Jan 1988, L.G. Clark & X. Londono 376 (COL, TULV, UPTC, US); Refugio del Ruiz, carratera hacia "El Silencio", 4310 m, 7 Oct 1978, Rangel et al. 1732 (COL); Cordillera Central, camino entre Buenos Aires y la Laguna Verde, 4020-4350 m, 9 Oct 1978, Rangel et al. 1791 (COL); Paramo entre Termales y Nevado Libano, 4620 m, 8 Dec 1958, Barclay & Juajibioy 6313 (COL); Risaralda: Mun. Pereira, below Nevado El Cisne, way from Casa del Cisne to Laguna Otiin, km 5,4100 m, 19 Sep 1999, D. Stancik 3378 (COL, PRC); Volcan de Santa Rosa, vertiente oriental, 4500 m, 20 Feb 1980, Jaramillo-Mejia et al. 5747 (COL); Entre Laguna El Ouin y Paramo de Santa Rosa, 4375 m, 4 Feb 1980, Jaramillo-Mejia et al. 6149 (COL); below Nevado de Santa Isabel, 4000 m, 20 Sep 1999, D. Stancik 3373B (COL, PRC); 4000-4100 m, 20 Sep \999,D. Stancik3374 (COL, PRC), 4300 m, 13 Jul 1980, Salamanca et al. 40 (US); junto a la Laguna del Beso, 4300 m, Festuca in South American Paramos 25 Nov 1946, J. Cuatrecasas 23201 (US); Parque Nacional Los Nevados, vertiente W del Nevado de Quindio, 4450 m, Salamanca etal. SSI43 (U). Tolima: Mun. Ibague, Nevado del Tolima, 04°38.5'N, 75°19.1'W, 4250 m, 8 Jun 2000, D. Stancik 3601, 3602, 3607 (COL, PRC); Mun. Santa Isabel, paso del Otiin a la Quebrada Africa, 3900 m, 6 Feb 1980, Jaramillo-Mejia etal. 6210 (COL); 4200-4300 m, Diaz-Piedrahita & Jaramillo-Mejia 1821 (COL). ECUADOR. Chimborazo: El Altar, N side, 4300 m, Sklendf & Kosteckova 1931 (AAU); Paramo de losAltares,0r40'S,78°24'W,4200m,3 Sep 1987, Ramsay & Smith 410 (K, QCNE); Volcan Chimborazo, lowerrefuge, 01°28'S, 78°50'W, 4800-4840 m, 11 May 1992, S. Laegaard 102797 (AAU, QCA, QCNE); S. Laegaard 102803 (AAU); 05 Mar 1988, 5 Laegaard & SA. Renvoize 70555 (AAU, K, QCA, QCNE); E slope, 4550 m, E. Asplund 7926 (S); Cerro Yuibug-Pailacajas, E side, 4100 m, Sklendf & Sklendovd 2990 (AAU); W side of Cerro Yanaurcu, 4300 m, Sklendf & Kosteckova 1853 (AAU). Imbabura: Nevado Cotacachi, SW side, 4400 m, Sklendf 6 Kosteckova 4300-4400 m, S. Laegaard 54518B (AAU, QCA); Mun. Cayambe, Volcan Cayambe, swamps below the Refugium, 00°00'N, 78°01'W, 4450 m, 20 Oct 2000, D. Stancik 4149 (PRC, QCA); rocky superparamo, 00°00'31 "S, 78°00'344"W, 4600 m, 20 Oct 2000, D. Stancik 4129, 4133, 4134, 4166 (AAU, PRC, QCA); D. Stancik 4122 (PRC, QCA); Mun. Urcuqui, Cerro Yanaurcu, rocks and pajonal, 00°29'N, 78°21"W, 4300 m, 15 Oct 2000, D. Stancik 4039 (AAU, PRC, QCA). Pichincha: Volcan Cayambe, 14000-14500 ft, Whymper 1330 (K); NE side, 15000 ft, Cazalet & Pennington 5749B (US); Cazalet & Pennington 5749 (K, NY); road to refuge, 4300-4400 m, 1 Mar 1988, S. Laegaard 70491 (AAU, K, QCA); SW side, 4600 m, Sklendf & Kosteckova 55-2 (US); Paso de la Virgen, 4400 m, 5 Laegaard 53869 (AAU, QCA, QCNE); Paramo de Guamani, 4200-4250 m, S. Laegaard55718 (AAU, QCA); 4200^1250 m, S. Laegaard 55696 (AAU); 00°20'S, 78°12'W, 4050 m, 2 Mar 1999, S. Laegaard etal. 19630 (AAU, QCA, QCNE); NE slope of Rucu Pichincha, 00°10'S, 78°34'W, 4500 m, Sklendf & Kosteckova 1903 (AAU); NE slope, 4300^1500 m, Sklendf & Kosteckova 1933 (AAU); W slope, 4400-4500 m, 6 Mar 1988, Molau & Eriksen 3298 (AAU, GB, QCA, QCNE); 4500 m, Sklendf 6 Kosteckova 9-1 (QCNE); 4350 m, E. Asplund 17334 (S); Guagua Pichincha, 00°12'S, 78°35'W, 4500-4600 m, S Laegaard et al. 102760 (AAU); Lloa-Guagua Pichincha road, km 10, 00°13'S, 109 78°35'W, 4170 m, S Laegaard et al. 102725 (AAU); Pichincha, 4700 m, E. Asplund 8594 (NY, QCA); 4100-4500 m, A.S. Hitchcock 21058 (NY, US); Sodiro s.n. (QPLS); Volcan Antisana, N side, 00°27'S, 78°10'W, 4700 m, S Laegaard 102889 (AAU); Nevado Cayambe, SW slope, 4400 m, Sklendf & Kosteckova 1893 (AAU); SW side, 4500^1700 m, Sklendf & Kosteckova 735 (AAU); 4220 m, 11 May 1990, P.M. Peterson 9086, E.J. Judziewicz & R.M. King (K, MO, QCA, QCNE, US); 4400 m, 01 Jul 1995, Sklendf & Kosteckova 49-19 (QCNE); 4600 m, 2 Jul 1995, Sklendf & Kosteckova 56-2 (QCNE); E slope of Illiniza Sur, 4300 m, Sklendf & Kosteckova 1863 (AAU); Cerro Saraurcu, W side, 4100-4200 m, Sklendf & Kosteckova 1836 (AAU); Mt. Corazon, E slope, 4500 m, E. Asplund 17504 (S); Mun. Pifo, Paramo Guamani, grassy paramo, 00°19'S, 78°12'W, 4300 m, 19 Jun 1999, D. Stancik 3031,3034 (AAU, PRC, QCA); 4000 m, D. Stancik 3048 (PRC, QCA). Tungurahua: Cordillera de Llanganatis, Paramo de Jaramillo, 01°10'S, 78°22'W, 4000-4250 m, S. Laegaard 53293 (AAU); S. Laegaard 53307 (AAU, QCA). 42. Festuca glyceriantha Pilg., Bot. Jahrb. Syst. 37: 516. 1906. (Figs. 52,54,93E & F). TYPE: Peru. Ancash: Yungay, Yanganuco, 4100 m, 15 Jun 1903, A. Weberbauer 3275 (holotype: B!; isotypes: BAA-1228 fragm. ex B!, US2875417!). Tussocked perennials with intravaginal innovations. Culms (30-)80-120 cm tall, erect, glabrous; nodes 1 or 2, basal. Leaf sheaths coriaceous, dark brown, glabrous, striate, old sheaths fibrous; ligules 0.5-1 mm long, apex acute, ciliate; blades 20-60 cm long, 2-5 mm wide, conduplicate or flat, abaxially glabrous, green or pale-green. Panicles 15-17 x 1.5-2 cm, contracted, lanceolate, discontinuous; branches scabrous. Spikelets 12-15 mm long, lanceolate, florets 4(-5); rachillas 2-2.5 mm long, glabrous, rarely scabrous; glumes 3.5-6.5 mm long, coriaceous, glabrous, violet, margins membranous; lower glumes 3.5-5 mm long, lanceolate, 1-nerved, apex acute; upper glumes (4.5-)5-6.5 mm long, lanceolate, 3-nerved, apex acute; lemmas 6-8.5 mm long, lanceolate, 5-nerved, coriaceous, violet, upper 1/3 with brown stripes, margin membranous, awned, the awn 0.52.5 mm long; paleas almost as long as the lemma, membranous, slightly scabrous; anthers 3-3.5 mm long; ovary apex glabrous. Caryopses not seen. Festuca in South American Paramos Figure 54. Festuca glyceriantha. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma with palea and rachilla. G. Lemma. H. Leaf blade cross-section. A-H, Vega 2012 (F). Festuca in South American Paramos Leaf blade anatomy.•Cross-sections with (7-) 11-15 vascular bundles and corresponding ribs above; sclerenchyma under abaxial epidermis discontinuous, extending to the vascular bundles, adaxial sclerenchyma present, extending often to the adaxial epidermis hairy; the hairs short. Observations.•Festuca glyceriantha is very distinct and easily recognized since it has dark brown, fibrous sheaths (often resembling bulbs) and brown stripes on the upper 1/2 of the lemma. Distribution and habitat.•Festuca glyceriantha is known from northern and central Peru where it occurs in alpine grass vegetation (jalca) between 330(M100(-4700) m. Additional specimens examined. PERU. Ancash: Prov. Huaylas, Huascaran Parque Nacional, Quebrada Los Cedros, Alpamayo trail, 4020^1700 m, 3 Nov 1985, Smith etal. 9939 (MO); 08°50'S, 77°44'W, 4020-4700 m, 3 Nov 1985, Smith 9942 (F, MO); Smith etal. 9944 (MO); Paron Vail., E of lake, 08°59'S, 77°38'W, 4200 m, 27 Sep 1985, Smith 11493 (MO); Prov. Huari, Huascaran Parque Nacional, quebrada Rurichinchay, 09°23 'S, 77°16'W, 3840-3870 m, 13 Jun 1986, Smith etal. 12630 (US); Cordillera Blanca, 28 km SE of San Luis on road towards Huari, 4230 m, 23 Mar 1997, P.M. Peterson 13875 & N. Refulio Rodriguez (US, USM); Prov. de Bolognesi, Caillarpacsa, arriba de Chuiquian, monte fluvial, 3900 m, 4 May 1952, Cerrate 1540 (US). Cajamarca: Prov. Celendin, lugar Challuayaco, sobre la carretera a Celendin, E del Paso de Cumullca, 3550 m, 5 Dec 1984, Vega et al. 3485 (F); Desvio a Huanico, sobre la ruta Cajamarca-Celendin, 3500 m, 4 Oct 1987, Vega 4371 (F); Sorochuco, arriba del Punre, 06°57'S, 78°16'W, 3340 m, 8 Nov 2001, Vega 10889 (F, MO); El Sendamal, entre la Encanada y Celendin, 3520 m, 7 Mar 1975, Vega et al. 1695 (F); Cerca al Centro Turistico Cumbe Mayo, 3460 m, 21 Apr 2001, Vega 10509 (F, MO); Cajamarca-Celendin road, 07°02'S, 78°14'W, 3450 m, 15 Jul 1983, Smith et al. 4278 (MO); Prov. Cajamarca, cerro Campanario, entre Cajamarca y Cooperativa Atahualpa, 3500 m, 8 Mar 1993, Vega 6616 (F); Prov. Cajamarca, Cerro Maqui-Maqui, al N de Cajamarca, ruta Shanta alta, 4120 m, 29 Jan 1994, Vega et al. 6694 (F); Cerro Campanario, en la SAIS Atahualpa, al NW de Cajamarca, 3500 m, 4 Nov 1994, Vega etal. 6974, 6975 (F); arboretum de CICAFOR, 3400 m, 16 May 1981, Vega et al. 2527 (F); Cerro Sexcemayo, al W de Cajamarca, 3430m,4Feb 1991, Vega5427(F); 3500m,20 Jun 111 1991, Vega et al. 5 744 (F); Jalca de Kumullca, entre La Encanada y Celendin, 3600 m, 27 May 1977, Vega etal. 2012 (F); Coymolache ruta CajamarcaHualgalloc, 3850 m, 7 Jan 1977, Vega et al. 2061 (F); 16 km W of Central Plaza of Cajamarca up road towards Cumbemayo, 3440 m, 31 Mar 1997, P.M. Peterson 14011 & N. Refulio Rodriguez (MO, US, USM); Prov. Hualgayoc, Perlamayo, Tres Lagunas, subiendo hacia Tambillo, 3200-2850 m, 22 Jul 1986, Vega 4168 (F); arriba de la ciudad de Hualgayoc a 31 km de Bambamarca, 3500 m, 26 Mar 1985, Vega et al. 3797 (F); Paso de Coymolache, km 81 de la carretera Cajamarca-Bambamarca, al W de Hualgayoc, 3900 m, 13 Jun 1980, Vega 2263 (F); al W de Hualgayoc, 3 km sobre la carretera a Hualgayoc-Cajamarca, 3600 m, 15 Jun 2001, Vega 10764 (F, MO); Prov. San Miguel, 64 km N of Cajamarca and 3 km SW of El Cobro, on route 3N towards Bambamarca, 3530 m, 16 Mar 2000, P.M. Petersonl4925 & N. Refulio Rodriguez (MO, US, USM); Prov. San Pablo, 32 km NW of Cajamarca on road towards San Pablo via Porcon, 3470 m, 14 Mar 2000, P.M. Petersonl4855 & N. Refulio Rodriguez (MO, US, USM); 2 km SW of Hualgayoc on road towards El Cobro and Cajamarca, 3530 m, 17 Mar 2000, P.M. Peterson 14936 & N. Refulio Rodriguez (MO, US, USM); Prov. De Hualgayoc, cerca a Coymolache entre Hualgayoc y Cajamarca, jalca, 3800 m, 16 Aug 1952, R. Ferreyra 8566 (US). Junin: NW part of Junin, Goyllarisquisca, 4000 m, 7 Nov 1923, A.S Hitchcock 22328 (US). La Libertad: Prov. Santiago de Chuco, 23 km SW of Huamachuco on road towards Alto de Tamboras and Pampas, 3540 m, 29 Mar 1997, P.M. Peterson 13961 & N. Refulio Rodriguez (MO, US, USM); Santiago-Shoreyo road, 35 km from Santiago, 08°03'S, 78°18'W, 4120 m, 26 Aug 1982, Smith 2337 (MO); Los Quinuales (al N de Quiruvilca), ladera, 3775 m, 24 Mar 1994, Leiva & Leiva 1094 (MO); Carrion, Trujillo-Huamachuco road, 07°56'S, 78°10'W, 3700 m, 13 Feb 1983, Smith et al. 3312 (MO). Prov. Huamachuco, Munmalca, Hda. Cochabamba, 3200 m, 26 Jun 1958, Lopez & Sagdsteguii 2827 (US). Piura: Prov. Huancabamba, Chulucanitas bajo, 3150 m, 5 Mar 1980, Vega 5271 (F). 43. Festuca holubii Stancik, Folia Geobot. Phytotax. 39(1): 102, f. 1, 6-10. 2004. (Figs. 52, 55). TYPE: Ecuador. Loja, Cerro de Arcos W of road Manu-Zaruma, 03°34'S, 79°28'W, 3250-3600 m, moist paramo, 14 Sep 1999, S 112 Festuca in South American Paramos 2 mm 2 mm Figure 55. Festuca holubii. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Laegaard &Aguirre 20611 (PRC). Festuca in South American Paramos Laegaard & Aguirre 20611 (holotype: AAU!; isotypes: LOJA!, PRC!, QCA!, QCNE!). Tussocked perennials with intravaginal innovations. Culms, 50-60 cm tall, erect, finely scabrous; nodes 1, basal. Leaf sheaths membranous, brownish-gray, glabrous, striate; ligules 0.6-0.8 mm long, two-lobed, apex short-ciliate; blades 15-25 cm long, 0.8 mm wide, conduplicate to involute, finely abaxially scabrous, green, apex obtuse. Panicles 8-14 cm long, ca. 0.7 cm wide, contracted, narrow, sparsely branched; branches scabrous on ribs. Spikelets 9-10 mm long, lanceolate, florets 3 or 4; rachilla glabrous or sparsely hairy; glumes 4-5.5 mm long, lanceolate to oblong-lanceolate, membranous to coriaceous, keeled, purplish, glabrous, apex obtuse and scabrous; lower glumes ca. 4 mm long, 1-nerved; upper glumes ca. 5.5 mm long, 3-nerved; lemmas ca. 7 mm long, oblonglanceolate, 5-nerved, membranous to coriaceous, purplish-green, distally keeled, scabrous, awned, the awn 1-2 mm long; callus glabrous; paleas 5/6 as long as the lemma, lanceolate, membranous, scabrous only on keels; lodicules 0.9-1 mm long, oblong, bilobed; anthers ca. 1.3 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 3/5-3/4 as long as the grain, linear. Leaf blade anatomy.•Cross-sections with 7 vascular bundles and 5 ribs above; sclerenchyma discontinuous, only under abaxial epidermis; adaxial epidermis with numerous hairs, the hairs ca. 0.09 mm long. Observations.•Festuca holubii is morphologically similar to the Peruvian F. divergens. However, the former differs from the latter by having taller culms (50-60 versus 20-35 cm), longer panicles (8-14 versus 6-7 cm), shorter spikelets are (9-10 versus 12-14 mm), and shorter lemmas (ca. 7 versus 10-12 mm). Distribution and habitat.•Festuca holubii is known only from the type collection in southern Ecuador and it occurs in grass paramos between 3200-3600 m. 44. Festuca huamachucensis Infantes, Revista Ci. (Lima) 54: 107.1952. (Figs. 56,57). TYPE: Peru. La Libertad, Huamachua Pro v., 7 Aug 1951, J. Infantes Vera 3538 (holotype: herb. Infantes; isotype: US-2044946!). Tussocked perennials with intravaginal innovations. Culms 15-55 cm tall, erect, glabrous; nodes 113 1 or 2, basal. Leaf sheaths membranous, brown, glabrous, striate; ligules 0.5-1 mm long, apex truncate, ciliate; blades 5-20 cm long, 0.6-0.8 mm wide, conduplicate to involute, abaxially glabrous, green, sometimes slightly curved. Panicles 15-17 x 0.5-1 cm, narrow, contracted, with few branches and these only near base; branches sparsely scabrous. Spikelets 8.5-10 mm long, lanceolate, florets 3 or 4, probably cleistogamous; rachillas 0.5-0.7 mm long, glabrous or sparsely scabrous; glumes 4-6.5 mm long, lanceolate, membranous, glabrous, apex acute; lower glumes 4-4.5 mm long, 1-nerved; upper glumes 5.5-6.5 mm long, 3-nerved; lemmas, 7-7.5 mm long, lanceolate, 5-nerved, membranous, glabrous, awned, the awn 4-6 mm long; paleas almost as long as the lemma, membranous; anthers 0.8-0.9 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 2/5 as long as the grain, linear. Leaf blade anatomy.•Cross-sections with 5(-7) vascular bundles, 3-5 ribs above; sclerenchyma under abaxial epidermis discontinuous, forming small fascicles, adaxial sclerenchyma absent or exceptionally forming small fascicles; adaxial epidermis with short scattered hairs. Distribution and habitat.•Festuca huamachucensis is endemic to northern Peru where it has been found growing on wet alpine grasslands (jalca). Additional specimens examined. PERU. Ancash: Prov. Recuay, Huascaran Parque National, quebrada Queshque, lateral valley toward Rio Pachacoto, 09°50S', 77°18'W, 4500^1600 m, 18 Mar 1986, Smith etal. 11851 (MO). Cajamarca: Prov. Cajamarca, 52 km N of Cajamarca, on Hwy 3N towards Bambamarca, and ca. 1 km W on a small road "Carretera Lagunas Yanacanchilla", 3800-3900 m, 16 Mar 2000, P.M. Petersonl4901 & N. Refulio Rodriguez (MO, US); Laguna MaquiMaqui, pajonal de jalca, 3910 m, 4 Oct 1994, Vega et al. 6955 (F); Cerro Piedras Gachas, entre Yanacocha y Llaucan, 3900 m, 23 Apr 1994, Vega et al. 6994 (F); cima de cerro con Piedras grandes, 3900 m, 24 Apr 1994, Vega et al. 7041 (F); cima del Cerro Maqui-Maqui, ladera arenosa, 4150 m, 20 Mar 1994, Vega et al. 6914 (F); Pampa Larga, al N de la explotacion minera Yanacocha, ladera de jalca graminosa, 3900 m, 14 May 1994, Vega 7124 (F); Jalca de Cumullca, lado occidental, 3600 m, 31 May 1984, Vega et al. 3494 (F); ruta a Guagal, km 10, 3600 m, 27 May 1977, Vega et al. 2018 (F); Ladera de jalca, 3900 m, 14 May 1994, Vega et al. 7128 (F); Prov. Celendin, desvio Huanico, 114 Festuca in South American Paramos 2 mm 2 mm >*/AFigure 56. Festuca huamachucensis. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf blade cross-section. A-G, Vega 6994 (F). sobre la ruta Cajamarca-Celendin, 3500 m, 4 Oct 1987, Vega 4378 (F); 07°02'31"S, 78°15'33"W, 3700 m, 26 Aug 2004, Sklendf & Zapata Cruz 8705 (PRC); Prov. Hualgayoc. Desvio de la carretera Coymolache-Chugur, 3750 m, 30 Apr 1994, Vega 7111 (F). La Libertad: Prov. Santiago de Chuco, 42 km S of Huamachuco on road towards Pampas, 4000 m, 29 Mar 1997, P.M. Peterson 13951 & N. Refulio Rodriguez (K, MO, US, USM); Prov. Carrion, al pie del Nevado de Huaylillas, jalca, Festuca in South American Paramos 07°52'S, 78°01 'W, 3840 m, 21 May 2001, Zapata 16506, 16508 (F). 45. Festuca imbaburensis Stancik, Folia Geobot. Phytotax. 39(1): 100, f. 3, 1-5. 2004. (Figs. 57, 58, 94A & B). TYPE: Ecuador. Imbabura, Mun. Urcuqui, road to Cerro Yanaurcu, 00°28'29"N, 78°20'04"W, grass paramo with swampy patches, 4150 m, 15 Oct 2000, D. Stancik 4098 (holotype: PRC!; isotypes: AAU!, QCA!). Tussocked perennials with intravaginal innovations. Culms (15-)20-50 cm tall, erect, finely scabrous; nodes 1, basal. Leaf sheaths membranous to coriaceous, stramineous, glabrous; ligules 0.5-0.7 mm long, membranous, apex truncate, short-ciliate; blades 9-20 cm long, 0.5-0.6 mm wide, conduplicate to involute, somewhat rigid, green, glaucous, abaxially glabrous, apex obtuse. Panicles 6-12 x ca. 0.5 cm, contracted, with few scabrous branches, each with a few spikelets. Spikelets 8-10 mm long, lanceolate, florets 2 or 3; rachilla glabrous; glumes 3-4.2 mm long, keeled on back, membranous, dark purple, glabrous or upper 1/4 sparsely papillose, apex obtuse; lower glumes 3-3.5 mm long, oblong, 1-nerved; upper glumes 4-4.2 mm long, ovate, 3-nerved; lemmas 5.5-6(-7) mm long, lanceolate, 5-nerved, membranous to coriaceous, dark purple, upper 1/4 scabrous, awned, the awn 0.5-0.7 mm long; callus glabrous; paleas shorter than the lemma, lanceolate, membranous, upper 1/4 and along keels densely scabrous to pilose; lodicules ovate, markedly bilobed; anthers 1-1.5 mm long; ovary apex glabrous. Caryopses not observed. Leaf blade anatomy.•Cross-sections with 5-7 vascular bundles and 3-5 ribs above; sclerenchyma under abaxial epidermis discontinuous, in small fascicles, adaxial sclerenchyma absent; 0 -85 -80 -75 115 -70 300 -65 600 Kilometers \_ "T" -60 Figure 57. Distribution of Festuca huamachucensis (•), F. imbaburensis (k.),F nereidaensis (•), and F. sanctae-martae (•). -10 116 Festuca in South American Paramos Figure 58. Festuca imbaburensis. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf blade cross-section. A-G, Stancik 3899 (PRC). Festuca in South American Paramos adaxial epidermis with scattered hairs, the hairs 0.09 mm long. Observations.•Festuca imbaburensis is morphologically similar to F glumosa but differs by having glabrous leaf blades (versus scabrous), narrower blades (0.5-0.6 versus 0.8-1.4 mm), shorter ligules (0.5-0.7 versus 0.8-1.2 mm), glabrous rachillas (versus pilose), and shorter lemmas (5.5-6(-7) versus 7-8 mm). Distribution and habitat.• Festuca imbaburensis ranges from central Ecuador to southern Colombia. It occurs in swampy patches associated with grass paramos and superparamos between 3400^1200 m. Additional specimens examined. COLOMBIA. Nariflo: Mun. Cumbal, Volcan Nevado de Cumbal, N-NE slopes, 3700 m, 9 Mar 1999, D. Stancik 2749 (COL, PRC, US); Mun. Tuquerres, Volcan Azufral, road from vereda San Roque alto to Laguna Verde, km 4,3700 m, 9 Mar 1999, D. Stancik 2783 (COL, PRC); road from vereda San Roque alto to Laguna Verde, km 5, 3800 m, 9 Mar 1999, D. Stancik 2786 (AAU, COL, PRC). ECUADOR. Azuay: Parque Nacional Cajas, trail from Molleturo to Paraguillas, 02°49'S, 79°15'W, 4200-4300 m, S. Laegaard 102666 (AAU); Mun. Cuenca, Parque Nacional Cajas, Cerro Amarrillo, 2°45'S, 79°15'W, 4300^1400 m, 13 Jul 1997, D. Stancik2056 (PRC, QCA); Cajas Parque Nacional, E flanks and summit area of Cerro Amarillo, 445 lm, lower superparamo with Loricaria sp., Azorella sp., Werneria sp., 79°15'W, 02°45'S, Sklenar 2596 (PRC, QCA). Bolivar: road Los Arenales-Salinas, km 4,01 °24' S, 78°55'W, 4300 m, S. Laegaard 55368 (AAU). Carchi: Paramo del Angel, sector Voladero, 00°38'N, 67°53'W, 3400-3800 m, Davalos 18 (US); Volcan Los Chiles, 00°49'N, 77°57'W, 3950-3980 m, S. Laegaard 101692 (AAU, QCA); Mun. Tulcan, volcan Chiles, SW slope, 00°48'N, 77°57'W, 4055m, 14 Jul 1999, D. Stancik 3228 (AAU, PRC, QCA, US, W); D. Stancik 3235,3267 (AAU, PRC, QCA). Chimborazo: El Altar, N side of Volcan, 4200^1400 m, Sklenar & Kosteckovd 956 (AAU); 4200 m, Sklenar & Kosteckovd 1929 (AAU); Sklenar & Kosteckovd 1031 (US); Collanes valley, Paramo de Los Altares, 4000 m, Ramsay & Smith 343 (QCA, QCNE); Alao-Paramo de Laguna Negra, 4100 m, Barclay & Juajibioy 8763 (MO, US); E side of Volcan Chimborazo, 4300 m, Sklenar & Kosteckovd 2168 (AAU); km 8.5 E of Guardiana Alao above Rio Alao, 3350-3550 m, P.M. Peterson 117 9197, E.J. Judziewicz, R.M. King & P.M. Jorgensen (MO, US); E of Licto, 4200 m, 30 Oct 1987, Ramsay 1035 (K, QCA, QCNE). Cotopaxi: road San Miguel (Salcedo)-Puerto Nuevo, 29 km from San Miguel, 3950-4050 m, B. Ollgaard & Balslev 9921 (AAU, F, MO, NY, S); Paramo de Cotopaxi, T. de Vries s.n. (AAU); Parque Nacional Cotopaxi, 4220 m, 21 Jun 1999, D. Stancik 3098 (AAU, PRC, QCA); Mun. Lasso, Parque Nacional Cotopaxi, NE side of volcano, 00° 39.3'S, 78°25'W, 4220 m, 26 Jun 1999, D. Stancik 3101 (AAU, PRC, QCA). Imbabura: Paramo de Mariano Acosta, km 25 on the road Yahuarcocha-Mariano Acosta, 00°20'S, 78°00'W, 3650-3750 m, S. Laegaard 101194 (AAU); Mun. Urcuqui, road to Cerro Yanaurcu, S slope, 00°28'56"N, 78°20'28.5"W, 4300 m, 15 Oct 2000, D. Stancik 4081 (AAU, PRC, QCA); D. Stancik 4090 (AAU, PRC, QCA, US); D. Stancik 4091 (AAU, PRC, QCA); D. Stancik 4092B, 4098 (PRC, QCA). Pichincha: road Pifo-Pintag, in valley 2.5 hours horseride above Inga Monserat, 00°19'S, 78°17'W, 3950 m, S. Laegaard 102262 (AAU); along road El Chaup-Pastocalle, 00°41 'S, 78°39' W, 3200-3700 m, S. Laegaard 54548 (MO); road Quito-Paramo Guamani, 00°17'S, 78°09'W, B. 0llgaard & Balslev 10073 (AAU, F, NY, MO, S); antenna N of pass, 00°17'S, 78°09'W, 4260xm, B. Ollgaard & Balslev 10100 (AAV,NY);00°lTS, 78°11 'W, 4250-4400 m, S Laegaard et al. 53859 (AAU, QCNE); 4050 m, S. Laegaard 101373 (AAU, QCA); peak ca. 6 km S of Paso de la Virgen, 00°21 'S, 78°13 'W, 4200-4250 m, S Laegaard 55698 (AAU); 4100 m, S. Laegaard 103105 (QCA); 3900 m, Ramsay et al. 153 (QCA, QCNE); N side of Volcan Antisana, ca. km 12 along road to Hacienda Antisana, 00°27'S, 78°10'W, 4400^1500 m, S. Laegaard 102871 (AAU); Mun. Pifo, Paramo de Guamani, 00° 20'S, 78°12'W, 4000 m, 19 Jun 1999, D. Stancik 3054B (PRC, QCA); D. Stancik 3054A (AAU, PRC, QCA); Superparamo vegetation on the S W side of Cotacachi, 0°21 'N, 78°21 'W, 4300^1400 m, 12 Aug 2004, Sklenar 8204 (PRC). Tungurahua: along trail from Mesa Tablon to Limpiopungu, S of Laguna Pisayambo, 01°07'S, 78°21'W, 4000-4050 m, S. Laegaard 19452 (AAU); Llanganatis, by Lake Aucacocha, 3700 m, Edwards s.n. (P); Mun. Pillaro, Las Llanganatis, 01°07'55.5" S, 78°20'40"W, 3900 m, 28 Sep 2000, D. Stancik 3898 (PRC, QCA); D. Stancik 3899 (AAU, PRC, QCA); 01°09'37.5"S, 78°14'51"W, 3500 m, D. Stancik 3990 (PRC, QCA). 118 Festuca in South American Paramos 1 mm Figure 59. Festuca monguensis. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik & Galvis2026 (PRC). Festuca in South American Paramos 46. Festuca monguensis Stancik, Darwiniana 41(1-4): 139, f. 8a-e. 2003. (Figs. 52, 59, 94C-F). TYPE: Colombia. Boyaca, Mun. Mongui, paramo de la Laguna La Colorada, 3550 m, 21 Jan 1999, D. Stancik & Galvis 2026 (holotype: PRC!; isotype: COL!, FMB!). Tussocked perennials with intravaginal innovations. Culms ca. 50 cm tall, erect, scabrous; nodes 2 on upper 1/2. Leaf sheaths coriaceous, stramineous, finely striate, glabrous; auricles absent; ligules 0.7-1 (-1.5) mm long, coriaceous, apex truncate, short-ciliate; blades 20-25 cm long, 0.6-0.8 mm wide, conduplicate to involute, abaxially scabrous, green, apex obtuse. Panicles 5-15 x 0.5-0.8 cm, contracted, lanceolate, with few branches; branches scabrous. Spikelets 9-11 mm long, lanceolate, florets 3 or 4; rachilla glabrous or with short, scattered hairs; glumes 4-5 mm long, lanceolate, coriaceous, purplish-green, scabrous near apex, margins membranous, apex acute; lower glumes ca. 4 mm, 1-nerved; upper glumes ca. 5 mm long, 3-nerved; lemmas 6-7 mm long, lanceolate, 5-nerved, membranous, purplish-green, papillose, scabrous near apex, awned, the awn 0.5-2.5 mm long; callus glabrous; paleas as long as the lemma, lanceolate, membranous, papillose, hairy along keel and near apex; lodicules 1.3 mm long, oblong, two-dentate; anthers 1.2-1.3 mm long; ovary apex glabrous. Caryopses not seen. Leaf blade anatomy.•Cross-sections usually with 7 vascular bundles, 5(-7) ribs above; sclerenchyma under abaxial epidermis discontinuous sometime extending to the vascular bundles, sclerenchyma absent adaxially; adaxial epidermis with scattered hairs, the hairs ca. 0.09 mm long. Observations.•Festuca monguensis differs from other species of F sect. Festuca by having short panicles on distinctive long culms. Distribution and habitat.•Festuca monguensis is endemic to Colombia and is known only from type locality in Cordillera Oriental (Boyaca). It is known from grass paramo dominated with Calamagrostis effusa (Kunth) Steud. between 3400-3600m. Additional specimens examined. COLOMBIA. Boyaca: Mun. Mongui, paramo de la Laguna La Colorada, 3550 m, 21 Jan 1999,£>. Stancik & Galvis 2025 (COL, PRC); 3450 m, 21 Jan 1999, D. Stancik & Galvis 2028 (COL, FMB, PRC). 119 47. Festuca nereidaensis Stancik, Darwiniana 41(1-4): 139, f. 8f-k. 2003. (Figs. 57, 60, 95A-D). TYPE: Colombia. Caldas, Mun. Manizales, Parque Nacional Los Nevados, road from Casa del Cisne to Rio Nereidas, km 5, 3800-4000 m, 18 Dec 1999,/). Stancik 3401B (holotype: PRC!; isotype: COL!). Tussocked perennials with intravaginal innovations. Culms 80-100 cm tall, erect, finely scabrous just below panicle; nodes 2 or 3 on upper 1/2. Leaf sheaths membranous to coriaceous, stramineous, glabrous; auricles absent; ligules 0.7-1 mm long, membranous, apex truncate, ciliate; blades 25-35 cm long, 1.2-1.5 mm wide, conduplicate, abaxially glabrous, green, apex acute. Panicles 15-20 x 1-1.5 cm, contracted, lanceolate, branches finely scabrous. Spikelets 11-12 mm long, lanceolate, florets 5; rachilla short, sparsely short-hairy; glumes 6-8 mm long, lanceolate, coriaceous, margins membranous, dark, glabrous, apex acute; lower glumes 6-6.5 mm long, 1 -nerved; upper glumes 7-8 mm long, 3-nerved; lemmas 7-7.5 mm long, lanceolate, 5-nerved, coriaceous, finely papillose, awned, apex pointed, the awn 0.5-0.7 mm long; callus glabrous; paleas almost as long as the lemma, lanceolate, membranous, papillose, upper 1/2 scabrous, apex hairy; lodicules 1.3 mm long, lanceolate, two-dentate; anthers 1.3-1.6 mm long; ovary apex glabrous. Caryopses not seen. Leaf blade anatomy.•Cross-sections with 10-12 vascular bundles and 9 ribs above; sclerenchyma under abaxial epidermis almost continuous, sometimes extending to the vascular bundles, adaxial sclerenchyma present but not extending to the vascular bundles; adaxial epidermis hairy, the hairs ca. 0.45 mm long. Distribution and habitat.•Festuca nereidaensis is endemic to Colombia and is known only from the type locality of Nevado del Ruiz in Cordillera Central. This species occurs in grass paramo zones between 3800^1000 m. 48. Festuca oroana Stancik, Folia Geobot. Phytotax. 39(1): 104, f. 2, 6-10. 2004. (Figs. 61, 64, 95E & F). TYPE: Ecuador. El Oro, along mule-track Tambillo-Manu, mountain forest, grazed and/or burned, dense tussock, 03°30'S, 79°32'W, 3200-3320 m, 21 Feb 1988, 21 Feb 1988, S. Laegaard 70309 (holotype: AAU!; isotypes: MO!, PRC!, QCA!, QCNE!). 120 Festuca in South American Paramos Figure 60. Festuca nereidaensis. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 3401B (PRC). Festuca in South American Paramos 121 Figure 61. Festuca oroana. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Laegaard 70309 (PRC). 122 Festuca in South American Paramos Tussocked perennials with intravaginal innovations. Culms ca. 100 cm tall, erect, scabrous; nodes 2(-3) on lower 1/2. Leaf sheaths membranous to coriaceous, lightly brownish-gray, finely scabrous, margins free; auricles absent; ligules 0.5-1 (-1.5) mm long, membranous, apex truncate, short-ciliate; blades 35-50 cm long, 1.1•1.8(•4) mm wide, conduplicate to flat, abaxially glabrous, green. Panicles ca. 20 cm long, ca. 25 cm wide, pyramidal, spreading, branches and pedicels densely minutely scabrous. Spikelets 9-10 mm long, lanceolate to oblong-lanceolate, florets 4 or 5(-6); rachilla densely hairy, the hairs short; glumes 2.4-3.9 mm long, membranous to coriaceous, green, apex acute; lower glumes 2.4-2.6 mm long, lanceolate, 1-nerved, glabrous; upper glumes 3.6-3.9 mm long, oblong to oblong-lanceolate, 3-nerved; lemmas 5.5-6(-6.5) mm long, lanceolate, 5-nerved, membranous, greenish, papillose or upper 1/3 finely scabrous, awned, the awn 0.5-1 mm long; callus hairy; paleas as long as the lemma, lanceolate, membranous, margins short-hairy, apex glabrous; lodicules oblong-lanceolate; anthers 2.7-2.8 mm long; ovary apex glabrous. Caryopses not seen. Leaf blade anatomy.•Cross-sections with about 17 vascular bundles and 9 ribs above; sclerenchyma under abaxial epidermis discontinuous, adaxial sclerenchyma extending to the vascular bundles forming girders; adaxial epidermis densely hairy, the hairs, 0.2-0.4 mm long. Observations.•In South America, there are no species morphologically similar to F. oroana. Perhaps Ffiebrigii is superficially similar but differs by having cylindrical panicles (versus pyramidal in F. oroana), longer glumes and lemmas (lower glume ca. 4 versus 2.4-2.6 mm; upper glumes 5-5.5 versus 3.6-3.9 mm; lemmas 6-7 versus 5.5-6 mm). Distribution and habitat.•Festuca oroana is known only from type locality in Southern Ecuador. It occurs in Andean mountain forests at an altitude of about 3200 m. 49. Festuca parciflora Swallen, Contr. U.S. Natl. Herb. 29(6): 255. 1949. (Figs. 62, 64). Ecuador. Azuay, paramo, in vicinity of Toreador, between Molleturo and Quinoas, 3810-3930 m, 15 Jun 1943, J. A. Steyermark 53092 (holotype: US-1911635!). TYPE: Tussocked, loosely caespitose or shortrhizomatous perennials with intravaginal innovations. Culms 28-60 cm tall, erect, scabrous or glabrous; nodes 1, basal. Leaf sheaths membranous to coriaceous, brownish-gray, glabrous, inconspicuously striate; ligules 0.8-1 mm long, membranous to coriaceous, apex truncate, short-ciliate; blades 13-23 cm long, 0.4-0.6 mm wide, conduplicate to involute, slightly abaxially scabrous, glaucous, apex obtuse. Panicles 3-15 cm long, ca. 0.5 cm wide, contracted, narrow with few erect branches; branches scabrous on ribs. Spikelets 8.5-9 mm long, lanceolate, florets 2 or 3(-4); rachilla with scattered hairs; glumes 3-5 mm long, keeled in upper part, membranous, purplish-white, upper 1/4 sparsely tuberculate; lower glumes 3-3.5 mm long, lanceolate, 1-nerved; upper glumes 4-5 mm long, ovate, 3-nerved; lemmas 6-7 mm long, lanceolate, 5-nerved, membranous, keeled, purplish-green, upper 1/3 scabrous, awned, the awn 0.5-1.5 mm long; callus with scattered hairs; paleas as long as the lemma, lanceolate, membranous, upper part and along keels densely scabrous to pilose; lodicules lanceolate; anthers 0.8-1.1(-1.4) mm long; ovary apex glabrous. Caryopses lanceolate; hilum 4/5 as long as the grain, linear. Leaf blade anatomy.•Cross-sections with 5 vascular bundles and 3 ribs above; sclerenchyma discontinuous under abaxial epidermis, adaxial sclerenchyma absent; adaxial epidermis with scattered hairs, the hairs 0.09 mm long. Observations.•This species is morphologically similar to F glumosa, F imbaburensis, F carchiense, and F sumapana but differs by having slender, scabrous leaf blades, smaller spikelets, and smaller floral parts. Distribution and habitat.•Festuca parciflora is endemic to southern Ecuador (Azuay, Loja) and is known from the grass paramo zone between 3600^1200 m. KEY TO THE SUBSPECIES OF FESTUCA PARCIFLORA la. Culms 40-50 cm tall, scabrous; plants tussocked; panicles 7•11(•15) cm tall 49a. F. parciflora subsp. parciflora lb. Culms ca. 25 cm tall, glabrous; plants short-rhizomatous to loosely caespitose; panicles 3-7 cm long 49b. F. parciflora subsp. loxana Festuca in South American Paramos 123 2 mm Figure 62. Festuca parciflora subsp. parciflora. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf blade cross-section. A-G Stancik 3846 (PRC). 124 Festuca in South American Paramos 49a. Festuca parciflora subsp. parciflora (Fig. 62). Tussocked perennials. Culms 40-50 cm tall, scabrous; panicles 7-11 (-15) cm tall. Additional specimens examined. ECUADOR. Azuay: Parque Nacional Cajas, N side of Laguna Luspa, 02°48'S, 79°15'W, 3850 m, 1 Sep 2000, D. Stancik 3845 (AAU, PRC); D. Stancik 3846 (AAU, PRC, US, W); Mun. Cuenca, Parque Nacional Cajas, between El Refugio and Lagoon Taglacocha-Tres Cruces, 02°47'S, 79°13'W, 39004000 m, 1 Sep 2000, D. Stancik 3859 (AAU, PRC); N side of Lagoon Taglacocha, 4100^1200 m, 1 Sep 2000, D. Stancik 3873 (AAU, PRC, US). 49b. Festuca parciflora subsp. loxana Stancik, Folia Geobot. Phytotax. 39(1): 107. 2004. TYPE: Ecuador. Loja, Cerro de Arcos W of road Manu-Zaruma, 03°34'S, 79°28'W, 3250-3600 m, 14 Sep 1999, S Laegaard & Aguirre 20608 (holotype: AAU!; isotypes: LOJA!, QCA!). Rhizomatous or loosely cespitose perennials. Culms ca. 25 cm tall, glabrous. Panicles 3-7 cm tall. Additional specimens examined. ECUADOR. Loja: Surroundings of Laguna Chuquiraga E of Amaluza, 04°37'S, 79°22'W, 3300 m, S Laegaard etal. 19285 (AAU). 50. Festuca renvoizei Stancik, Novon 17(1): 100-104.2007. (Figs. 63,64). TYPE: Peru. Cajamarca, Las Lagunas, 50 km from Cajamarca on road to Bambamarca, heavily grazed and burned grass paramo with crystalline rock outcrops and numerous lagoons of various size, growing in very wet area beside lagoon, 4000 m, 29 Mar 1988, S.A. Renvoize & S. Laegaard 5031 (holotype: AAU!; isotypes: CPUN!, K!). Tussocked perennials with intravaginal innovations. Culms 40-50 cm tall, erect, scabrous; nodes 1, basal. Leaf sheaths membranous, brownish-gray, abaxially glabrous, margins free; auricles absent; ligules 1.2-1.5 mm long, membranous, shortciliate; blades 20-30 cm long, 0.5-0.7 mm wide, conduplicate to involute, mostly glabrous, green, apex obtuse. Panicles 11-17 cm long, 0.5-0.7 cm wide, contracted, slender; branches scabrous throughout. Spikelets 9-10.5 mm long, lanceolate, florets 2 or 3; rachilla glabrous; glumes 3.5-5.5 mm long, membranous, dark purple, mostly glabrous, apex acute; lower glumes 3.5^4.5 mm long, lanceolate, 1-nerved; upper glumes 5-5.5 mm long, oblong-lanceolate, 3-nerved; lemmas 7-7.5 mm long, lanceolate, 5-nerved, membranous, dark purple, scabrous distally, awned, the awn 0.5-1 mm long; callus glabrous; paleas 4/5 as long as the lemma, membranous, upper 1/3 and along keels hairy; lodicules ca. 0.8 mm long, lanceolate; anthers 1-1.2 mm long; ovary apex glabrous. Caryopses not observed. Leaf blade anatomy.•Cross-sections with 5-8 vascular bundles, 5 ribs above; sclerenchyma under abaxial epidermis discontinuous and small, adaxially absent; adaxial epidermis with scattered hairs, the hairs ca. 0.1 mm long. Distribution and habitat.•Festuca renvoizei is known only from Department Cajamarca in northern Peru from grass paramos in very wet areas near lagoons at 4000 m. Additional specimens examined. Peru. Cajamarca: Parte baja del Cerro Shillas Negras y Laguna Totora, 3920 m, 23 Apr 1994, Vega et al. 7011 (F, CPUN). 51. Festuca sanctae-martae Stancik, Preslia 75(4): 343-345, f. 2. 2004. (Figs. 57, 65, 96A-D). TYPE: Colombia. Magdalena, Sierra Nevada de Santa Marta, SE slope, Hoya del Rio Donachui, Laguna de Calocribe (E of Meollaca), paramo, 3600-3700 m, 30 Sep 1959, J. Cuatrecasas & Castaneda 24532 (holotype: COL!; isotype: US!). Tussocked perennials with intravaginal innovations. Culms 60-80 cm tall, erect, glabrous or finely scabrous under panicle; nodes 1, basal and 1 or 2 shorter leaves. Leaf sheaths membranous to coriaceous, stramineous, glabrous, finely striate; auricles absent; ligules 1-1.6 mm long, membranous to coriaceous, apex truncate; blades 25-35 cm long, (0.8-)2-3.5 mm wide, conduplicate or flat, abaxially glabrous. Panicles 10-16 x 1.5-2 cm, contracted, elongate. Spikelets 10-13 mm long, lanceolate, florets 2 or 3; rachilla pubescent; glumes 8.5-10.5 mm long, almost as long as the spikelet, lanceolate, membranous, white to purplish-white, scabrous along midnerve, apex acute; lower glumes 8.5-9.5 mm long, 1-nerved; upper glumes 8.5-10.5 m long, 3-nerved; lemmas 9-10 mm long, lanceolate, 5-nerved, membranous, awnless or short-awned, the awn 0.5-1 mm long; callus Festuca in South American Paramos 125 Figure 63. Festuca renvoizei. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Renvoize & Laegaard 5031 (AAU). Festuca in South American Paramos 126 70 it 4-80 -75 -70 Figure 64. Distribution of Festuca vaginalis (•), F. parciftora (A), F. oroana (•), and F. renvoizei (•). glabrous or sparsely hairy; paleas 3/4 as long as the lemma, papillose, upper 1/3 scabrous; lodicules 1-1.4 mm long, lanceolate; anthers 0.8-1.1 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 3/4 as long as the grain, linear. Leaf blade anatomy.•Cross-sections with (8•)10•11 vascular bundles, with 5-7 ribs above; sclerenchyma under both abaxial and adaxial epidermis, discontinuous, sclerenchyma girders absent; bulliform cells not observed; adaxial epidermis with scattered hairs, the hairs ca. 0.02 mm long. Observations.•Festuca sanctae-martae is morphologically similar to F. glumosa and F cocuyana. Festuca sanctae-martae and F cocuyana differ from F glumosa by having conduplicate (versus involute) leaf blades with 8-11 (versus 5-7) vascular bundles, lower glumes 7.5-9.5 mm long (versus 4.5-6 mm), upper glumes 7.5-10.5 mm long (versus 6-6.5 mm), and lemmas 8-10 mm long (versus 7-8 mm). Festuca cocuyana differs from F sanctae-martae by having shorter culms (20-50 versus 60-80 cm), shorter leaf blades (15-20 versus 25-35 cm), and shorter lower glumes (7.5-8 versus 8.5-9.5 mm). Distribution and habitat.•Festuca sanctaemartae is endemic to the Sierra Nevada de Santa Marta in northern Colombia. This species is known from the grass paramos dominated with Calamagrostis effusa and shrubby matorral zones between 3600^1300 m. Additional specimens examined. COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, valley descending SW from Picos Reina and Ojeda, around Lagoons Naboba, Mamito, and Mamo, 4200-4300 m, J. Cuatrecasas & Castaneda 24563 (COL, US). 52. Festuca subulifolia Benth., PI. Hartw. 262. 1846. (Figs. 66,67,96E-F). Festuca tolucensis Festuca in South American Paramos 127 Figure 65. Festuca sanctae-martae. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Cuatrecasas & Castaneda 24532 (COL). 128 Festuca in South American Paramos 1 mm Figure 66. Festuca subulifolia. A. Stylized growth form. B. Habit. C. Inflorescence with proliferating spikelets. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-B, D-I, Stancik 3013 (PRC); C, Stancik 3336 (PRC). Festuca in South American Paramos -85 -80 -75 -70 129 -60 -65 * ^ ""A. PANAMA 10 10 /r *Y-\% ^ Jj VENEZUELA • COLOMBIA \ / \ BRAZIL f^ECUAlXm *x \ ikrn iti \4 N ^•1L. -10 -85 -80 -75 -70 0 ' \ 300 600 Kilometers A. -10 ~ \ -65 -60 Figure 67. Distribution of Festuca tolucensis (•), F. subulifolia (A), and F. sumapana (•). var. subulifolia (Benth.) St.-Yves, Candollea 2: 302, 316. 1925. TYPE: Ecuador. Pichincha, Hacienda de Antisana, Hartweg 1455 (lectotype: K!, designated by Alexeev, Novosti Sist. Vyss.Rast.23:18.1986;isolectotypes:B!,NY exK!,P!,W!). Festuca breviaristata Pilg., Bot. Jahrb. Syst. 25(5): 717-718. 1898. TYPE: Ecuador. Pichincha, Mount Puntas, 4400 m, Oct-Nov 1971, A. Stiibel207 (holotype: B!; isotypes: US-2875385 fragm. ex B!, S!). Festuca cajamarcae Pilg., Bot. Jahrb. Syst. 37: 513. 1906. Festuca distichovaginata var. cajamarcae (Pilg.) St.-Yves, Candollea 3 :222. 1927. TYPE: Peru. Cajamarca, Pas Coymolache, 4000-4100 m, May 1904, A Weberbauer 3974 (holotype: B!; isotypes: BAA-1171 fragm.!, US-2875389!). Festuca ferreyrae Tovar, Publ. Mus. Hist. Nat. Javier Prado, Ser. B, Bot. 32: 8. 1984. TYPE: Peru. Amazonas, Prov. Chachapoyas, Leimebamba, en pajonal, 17 Apr 1964, R. Ferreyra 15512 (holotype: US-2565588!). Dense tussocks (25-)50-80 cm tall; innovations intravaginal; culms erect, finely scabrid or glabrous, with one basal node and 2 leaves. Sheaths coriaceous, firm, stramineous (or greyish), glabrous (or finely scabrid); ligule coriaceous-membranous, two-lobed, shortly ciliate, (0.5•)1•2.5 mm long; leaf blades linear, conduplicate, pungent, 10-60 cm long, 0.4-0.8 mm in diameter, rough, green to glaucous. Panicle usually contracted, dense, lanceolate to oblong, some times spreading, ovate, (6-) 12-3 5 long, 1•2(•10) cm wide; branches finely scabrid. Spikelets 6.5-9 mm long, ovate, with 3-5 florets; rachilla covered (densely) by hairs; glumes coriaceous-membranous, acute, green or purplish-green, upper third (or all) scabrous, nerves prominent; lower glume lanceolate, (3-)3.5-4.5(-5.5) mm long, 130 Festuca in South American Paramos 1-nerved; upper glume oblong, 4-5.5(-6) mm long, 3-nerved; lemmas 5-6.5(-8) mm long, 5-nerved, coriaceous-membranous (on margins), oblonglanceolate, purplish-green, densely scabrid to rough on all surfaces, shortly two-dentate, with awn 0.5-2.5 mm long; callus glabrous; palea membranous, lanceolate, two-carinate, at least upper third and along keels scabrid to hairy, two-dentate, as long as the lemma (or longer); lodicules obovate, two-dentate, about 1.1 mm long; stamens 3, anthers (2. l-)2.4•2.8(3.2) mm; ovary apex glabrous. Caryopsis oblong-lanceolate; hilum linear, 4/5 of total. Leaf blade cross section typically with 7 vascular bundles and 5 ribs above; sclerenchyma under abaxial epidermis continuos with all or almost all vascular bundles, adaxial sclerenchyma present at 3 vascular bundles and forming girders, (about 30% of studied specimens with 9-13 vascular bundles forming 7-11 ribs and with 3-6 girders); abaxial epidermis densely covered by prickles, hairs more or less dense on adaxial epidermis, 0.1-0.2 mm long. Leaf blade anatomy.•Cross-sections typically with 7 vascular bundles and 5 ribs above; sclerenchyma under abaxial epidermis continuous extending to (almost) all vascular bundles; adaxial sclerenchyma present and extending to 3 vascular bundles forming girders, ca. 30% of studied specimens with 9-13 vascular bundles forming 7-11 ribs and with 3-6 girders); abaxial epidermis densely covered by prickles, densely hairy on adaxial epidermis, the hairs 0.1-0.2 mm long. Observations.•Festuca subulifolia is morphologically similar to F. tolucensis, however F. subulifolia differs by having large girders in leaf blade cross sections and shorter truncate ligules. Festuca subulifolia is a highly variable species that is often a dominant or co-dominant in the grass paramo zone. The variation in form and size of the panicles and floral parts cannot be taxonomically interpreted easily and it is probably a result of the diversity of natural conditions and of paramo management, i.e., burning and grazing. Characteristics, such as two-dentate lemmas with a relatively long awn and finely scabrous leaf blades, are fairly stable with not much variation. After comparing the types of Peruvian material of F cajamarcae and F. ferreyrae, with that of F subulifolia, it was concluded that these types represent slightly different forms of the same species. Therefore the former two names are here treated as synonyms. Distribution and habitat.•Festuca subulifolia is known from Peru, Ecuador, and Colombia. In Colombia Festuca subulifolia is known only in southern part of Cordillera Central (Narino, Cauca). It is often a dominant species of grass paramos between 2900-3700 m. This species is known from various plant communities such as: Agrostis cf. haenkeana (Cuatrecasas 1934), Calamagrostio effusae-Calamagrostietum macrophyllae (Duque & Rangel 1989), Aciachne acicularis-Calamagrostis intermedia (Verweij 1995), Ugno myricoidisEspeletietum hartwegianae (Rangel & Ariza 2000), Espeletio hartwegianae-Calamagrostietum effusae (Salamanca 1991), and Calamagrostis effusaeEspeletia hartwegiana subsp. centroandina (Rangel & Franco 1985). Additional specimens examined. COLOMBIA. Cauca: Mun. Popayan, Parque Nacional Purace, Volcan Purace, 02°20.07'N, 76°23.65'W, 42004300 m, 6 Jul 2000, D. Stancik 3609 (COL, PRC); Pilimbala, 02°22.1'N, 76°24.06'W, 3350 m, 6 Jul 2000, D. Stancik 3608 (COL, PRC), 3450 m, 2 Apr 1939, Kjell von Sneidern 2144 (K, US); Macizo Colombiano, Valle de las Papas, alrededores de Valencia, 3500-3700 m, 15 Sep 1981, Idrobo etal. 3834 (COL, P, US); Hda. Los Andes, 3150 m, 17 Sep 1958, Barclay 5865 (COL); alrededores de la Laguna de Cusiyaco, 3360 m, 7 Oct 1958, Barclay & Juajibioy 5942 (COL, MO); 3360 m, 7 Oct 1958, Barclay & Juajibioy 5945 (COL, MO, US); 12 Oct 1958, Barclay & Juajibioy 6052 (COL, MO); Paramo del Purace, E slope, around Laguna San Rafael, 3320 m, 11 Oct 1961, J. Cuatrecasas & Willard 26292 (COL, US); 3300 m, 27 Dec 1988, Ortiz 1272 (COL); Laguna San Rafael, 3300 m, 6 Apr 1985, Wood 4806, 4807 (COL, K); Paramo El Hinchadera, 3700 m, 24 Jul 1943, J. Cuatrecasas 14694 (US, VALLE); Cabeceras del Rio Paez, Laguna del Paez, 3450 m, 4 Dec 1944, J. Cuatrecasas 19059 (US, MO, VALLE); Paramo de Moras, 9 Mar 1999, D. Stancik 2685 (COL, PRC); 30003500 m, Feb 1906, Pittier 1409 (US); Mt. Pan de Azucar, 3500-3700 m, 16 Jun 1922, F Pennell 7057 (US). Narino: El Encano, km 6 in direction to village Colon, 2900 m, 13 Mar 1999, D. Stancik 2860 (COL, PRC, PSO); 13 Mar 1999, D. Stancik 2856 (COL, PRC); Mun. Guachucal, vereda Quetanbu, Paramo Infiernillo, 3200 m, 9 Mar 1999, D. Stancik2613, 2618 (COL, PRC, PSO); 3100 m, 9 Mar 1999, D. Stancik 2617, 2619 (COL, PRC, PSO); D. Stancik 2631 (COL, PRC); Mun. Pasto, paramo Puerto frio, between villages Las Almas and Alisales, 2900 m, 14 Mar 1999, D. Stancik 2876 (COL, PRC, PSO); Mun. Pasto, Volcan de Festuca in South American Paramos Galeras, 3400 m, 16 Mar 1999, D. Stancik 2911 (COL, PRC, PSO); 3200 m, 16 Mar 1999, D. Stancik 2909 (COL, PRC, PSO); 9 Mar 1999, D. Stancik 2659 (COL, PRC, PSO); 3600 m, 16 Mar 1999, D. Stancik2918 (COL, PRC, PSO); 2900 m, 16 Mar 1999, D. Stancik 2910 (COL, PRC, PSO); 16 Mar 1999, D. Stancik 2908 (COL, PRC, PSO); Mun. Pasto, Morazurco, vereda San Francisco, 3100 m, 21 Mar 1999, D. Stancik 2962 (COL, PRC, PSO); 21 Mar 1999, D. Stancik 2963 (COL, PRC, PSO); Village Piedrancha, Paramo Infernillo, 3200 m, 9 Mar 1999, D. Stancik2659 (COL, PRC, PSO); 9 Mar 1999, D. Stancik 2656, 2657 (COL, PRC, PSO); D. Stancik 2658 (COL, PRC); Mun. Tuquerres, Volcan Azufral, vereda San Roque Alto, 3600 m, 9 Mar 1999, D. Stancik2764 (COL, PRC, PSO); D. Stancik 2766 (COL, PRC); 9 Mar 1999, D. Stancik 2802 (COL, PRC, PSO); 9 Mar 1999, D. Stancik 2789 (COL, PRC); E slope, 0P07'N, 77°40'W, 3100-3300 m, 15 Jun 1995, J. Luteyn et al. 12834 (COL, MO); 12 May 1989, J. Luteyn et al. 12842 (COL); Paramo Bordoncillo, 3250 m, 5 Mar 1963, Espinal 1035 (COL, PSO, US); Chiles, cerca de poblacion Chiles, 3200 m, 30 Oct 1955, Fernandez-Perez 2931 (COL); Paramo de Uillinsayaco, between Pasto y Sibundoy, 15 Jul 1957, Barclay 4529 (COL, MO); Volcan Galeras, 2900 m, 30 Oct 1980, Benavides 2660 (PSO); 3500 m, 21 Mar 1986, Benavides 6554 (PSO); 3400 m, 25 Aug 1964, Mora-Osejo 3195 (PSO); 3600 m, 5 Feb 1965, Mora-Osejo 3503 (PSO); Cumbal, 4000 m, 19 Mar 1941, Kjell von Sneider 362 (US); 19 Mar 1941, Kjell von Sneider 435 (US); 3700 m, 24 Mar 1941, Kjell von Sneider 415 (US); Mun. Pasto, Santuario Volcano Galeras, 2900-3600 m, 16 Mar 1999, D. Stancik 2914, 2917 (COL, PRC); Mun. Cumbal, Nevado del Cumbal, N-NE slope, from vereda Las Vuertas, 3700 m, 5 Sep 1999, D. Stancik 2748 (COL, PRC). ECUADOR. Azuay: Paramo SW of Cuenca, 02°55'S, 79°08'W, 3500 m, 1 Sep 1984, S. Laegaard 52837 (AAV, K, QCA, QCNE); Road Gualaceo-Sucua, 03°00'S, 78°40'W, 3450 m, 23 Oct 1984, S. Laegaard 53213 (AAU, QCA); along new road Cuenca-Saraguro, km 14 N of sideroad to Nabon, 03°12'S, 79°02'W, 3420 m, S Laegaard 102376 (AAU); 03°07'S, 79°19'W, 3250-3350 m, 22 Mar 1992, S. Laegaard 101856 (QCA, QCNE); Parque Nacional Cajas, road to Molleturo, km 38.4, 02°46'S, 79°14'W, 41104350 m, P.M, Jorgensen et al. 2099 (MO, PRC); P.M. Jorgensen 2074 (MO, PRC); Laguna Torreadora, 02°46'50"S, 79°13'40"W, 4100-4150 m, 131 2 Aug 1999, Police 6, 7, 8 (PRC); N side of Lagoon Taglococha, 02°47'S, 79°15'W, 4100-4200 m, 1 Sep 2000, D. Stancik 3847, 3848, 3850, 3872 (PRC, QCA); slopes near Laguna Luspa and Laguna Canutillos, 3700 m, 21 Apr 1990, P.M. Peterson 8865, C.R. Annable & M.E. Poston (K, MO). Totorococha-Mazan Valley, 02°53'S, 79°10'W, 3900 m, Ramsay & Smith 510 (K, QCA, QCNE), vicinity of Toreador, 3810-3930 m, J. Steyermark 53146 (F, NY, US); J. Steyermark 53182 (US); Paramo de Tinajillas, ca. 10 km S of Cumbe, 3300m,Harling 11255(GB);km 13.7 S of Cumbe, 3470 m, P.M. Peterson 8876, C.R. Annable & M.E. Poston (US); Paramo de Patococha, entre Gualaceo y Limon, 3450-3500 m, Barclay & Juajibioy 8641 (MO, US); 03°00'S, 78°38'W, 3400 m, S. Laegaard et al. 103067 (AAU, QCA, QCNE); km 45.4 W of Indanza on road to Cuenca, 3300 m, P.M. Peterson 8932, C.R. Annable & M.E. Poston (K, MO, QCA, QCNE, US); between el Refugio y Lagoon Taglacocha, 02°47'S, 79°13'W, 39004000 m, 1 Sep 2000, D. Stancik 3862, 3863, 3864, 3865, (PRC, QCA); Lagoon Luspa, S side, 02°48'S, 79°15'W, 3850 m, 1 Sep 2000, D. Stancik 3837 (PRC, QCA); Mun. Nabon, road Loja-Cuenca, Cerro Ungahipucera, 03°12'S, 79°02'W, 33003450 m, 30 Aug 2000,/). Stancik 3804, 3805, 3811, 3815 (PRC, QCA); Bolivar: pass on road Guaranda-Riobamba, NE of road, 01°35'S, 78°50'W, 4050^1150 m, 10 Jun 1990, S Laegaard 71739, 71741 (AAU, QCNE); km 27, 01°38'S, 78°51'W, 3900^1100 m, S Laegaard 55001 (AAU, QCA); highway Ambato-Guaranda km 55.4, 4050 m, P. M. Peterson 8983 & C.R. Annable (MO, QCA, QCNE, US); km 60.4,4090 m, P.M. Peterson 8985 & C.R. Annable (K, MO, QCA, QCNE, US). Canar: at the antennas of Culebrillas, ca. 17 km from Panamerican Hwy., 02°26'S, 78°57'W, 4000 m, 4 Feb 2000, S Laegaard 21016 (AAU, LOJA); road Canar-Chunchi, 02°24'S, 78°59'W, 3200 m, S. Laegaard 53834 (AAU, QCA, QCNE); nudo de Cordillera Oriental and Occidental, 3240 m, Barclay & Juajibioy 8325 (COL, US); km 11.4 NW of Tambo, S facing slope, 3220 m, P.M. Peterson 8846, C. R. Annable & M.E. Poston (QCNE). Carchi: Paramo del Angel, sector Los Voladores, 00°41'N, 77°53'W, 3700-3750 m, S. Laegaard 101264A (AAU, QCA); NE of El Angel, 3700-3900 m, S Laegaard 53125 (AAU, QCA, QCNE); 3600 m, Harlington 4067 (S); 3400-3800m,Davalos21, 28(\JS); 3200-3800m, S. Laegaard 55771 (AAU, QCA); 00°4LN, 132 Festuca in South American Paramos 78°54'W, Holm-Nielsen 5427 (AAU, COL, F, MO, S); 00°50'N, 77°55'W, 3800 m, Grignon 84189 (AAU); 3750-3850 m, Holm-Nielsen 5383 (F); 3800 m, Acosta-Solis 10538 (US); km 13 on road Las Juntas-El Angel, 00°43'N, 77°50'W, 3380 m, S. Laegaard 101726B (AAU, QCA, QCNE); 3300 m, E. Asplund 7055 (S); 3900 m, E. Asplund 17022 (S); km 4 W of Tufino, Monte Redondo area, 3475m, J. Luteyn & Luteyn 5713 (AAU, MO, NY, US); NE of road El Angel towards Tulcan, 3240 m, P.M. Peterson 9122, E.J. Judziewicz & R.M. King (K, MO, QCA, QCNE, US); 13 May 1990, P.M. Peterson 9127, E.J. Judziewicz & R.M. King (K, MO, QCA, QCNE, US); 3320 m, P.M. Peterson 9140, E.J. Judziewicz & R.M. King (K, MO, QCA, US); Nudo de Boliche-Voladero, 3900 m, Penland & Summers 889 (F); Road Tulcan-El Carmelo, 3300 m, Harling & Andersson 12538 (GB); Paramo del Angel, 30 Dec 1931, Benoist 4636 (P); Mun. Tulcan, volcan Chiles, around Laguna Verde, 00°48 'N, 77°56' W, 4000 m, 14 Jul 1999, D. Stancik 3240 (PRC, QCA). Chimborazo: Parque Nacional Sangay-comunidad Alao-Llactapamba, 01°58'S, 78°28'W, 3500 m, Ceron et al. 11813 (MO, QCNE); Altar, A peakValley of Collanes, 13000 ft, Whymper 1638 (K); Collanes valley, 01°40'S, 78°24'W, 4200 m, Ramsay & Smith 422 (K); Panamerica Highway Riobamba-Ambato, 01°30'S, 78°42'W, 3500 m, S. Laegaard 55401 (AAU, QCA); W of pass Alao-Huamboya, 01°48'S, 78°25'W, 3750-3800 m, S. Laegaard 55417 (AAU, QCA); E of Guardiana Alao, km 8.5 via Huamboya, 3350-3550 m, P.M. Peterson 9189, E.J. Judziewicz, R.M. King & P.M. Jorgensen (MO, US); Paramo de Chacheaco, E de Chunchi, 3800 m, Barclay & Juajibioy 8264 (MO, US); Daldal Valley E of Licto, km 10, 01°48'S, 78°32'W, 3850 m, Ramsay et al. 1057 (QCA); Mt. Chimborazo, E slope, 4250 m, E. Asplund 8402 (NY, QCA, S), 3650 m, E. Asplund 7880 (F, S); road to Pallatanga, km 10 E of Lago Colta, 3725 m, P.M. Peterson 9215, E.J. Judziewicz, R.M. King & P.M. Jorgensen (QCA, QCNE, US); along Whymper road, ca. 10 km S of Cruce de los Arenales, 01°30'S, 78°52'W, 4100 m, S. Laegaard 19148 (AAU, LOJA, QCNE); ChimborazoReserva faunistica, 01°30'S, 78°49'W, 4150 m, Ceron et al. 19819 (QAP); Pungala-comunidad Alao, 01°52'S, 78°30'W, Ceron 28957 (QAP); along Rio Alao, 01 °52'S, 78°30' W, 3200-3400 m, S. Laegaard 55304 (AAU, QCA); Paramo de Urbina, 3600 m, Acosta-Solis 21195, 21208, 21216 (US); 3600-4500 m, A.S. Hitchcock21944, 21951 (US); 3700 m, E. Asplund 7803 (S); 3600 m, A.S. Hitchcock 22030 (NY, US); road El Pocuy-El Arenal, 3800 m, Acosta-Solis 166989 (US); Chimborazo, 4100^1400 m, Rauh & Hirsch E321, E331 (US); W slope of Chimborazo, 01°30'S, 77°50'W, 4000 m, Ramsay & Smith 1016 (K); E side, 3480 m, Barclay & Juajibioy 8819 (COL, MO, US); lado A, 3500 m, Barclay & Juajibioy 8162 (COL, MO, US); Bosque Andino de Cubillin, 3300-3400 m, Acosta-Solis 7554 (US); paramo along the road to Guaranda, 4150 m, E. Asplund 8191 (S); road Pallatanga-Cajabanba, km 47 from Pallatanga, 3750-3800 m, B. Ollgaard & Balslev 8926 (AAU, MO, NY); B. Ollgaard & Balslev 8962 (F); road to lago Colta, km 9 NE of San Juan, 3600 m, 21 May 1990, P.M. Peterson 9245, E.J. Judziewicz, R.M. King & P.M. Jorgensen (K, MO, QCA, QCNE, US); Entre Culebrillas y Yanayacu, 3300-3450 m, Acosta-Solis 7614, 7623 (F, S, US); Mun. Guaranda, W side of the volcan Chimborazo, 01°28'S, 78°52'W, 4100 m, 5 Jul 1999, D. Stancik 3162, 3164, 3169, 3189, 3190, 3193 (PRC, QCA); NNW side of volcan, above vereda Rio Colorado, 4150 m, 2 Jul 1999, D. Stancik 3176 (PRC, QCA); Mun. Riobamba, Volcan Chimborazo, sector Cruce de los Arenales, 01°28'14.6"S, 78°54'06"W, 4300 m, 20 Sep 2000, D. Stancik 3705B, 3707, 3708, 3709, 3714, 3715, 3716, 5777 (PRC, QCA. Cotopaxi: Parque Nacional Cotopaxi, falda NNW, 4000 m, Ehrenburg 1, 27, 36 (QCA); E of Loma Ingapirca, 00°40'S, 78°30'W, 4000 m, 6 Nov 1982, Balslev et al. 3435, 3438 (QCA); Balslev et al. 3437 (MO, QCA, US); falda N de Cotopaxi, 4000 m, Balslev et al. 3675, 3733, 3734, 3738 (QCA); Cerro Sinfana, 00°40'S, 78°28'W, 15 Dec 1990, Ceron 12609 (QAP); railway station Cotopaxi, 3500 m, E. Asplund 16807 (S); 3400 m, E. Asplund 6471 (F, MO, S, US); 3500 m, E. Asplund 18286 (S); 3350 m, 13 Apr 1990, P.M. Peterson 8721 (K, MO, QCA, QCNE, US); 3550 m, E. Asplund 6369 (F, S); quebrada de Agualongo, 00°40'S, 78°30'W, 3000 m, 10 Jan 1994, Ceron 18088 (QAP); Laguna Limpiopungo, 3800 m, 14 Apr 1990, P.M. Peterson 8744, C.R. Annable & M.E. Poston (K, MO, QCA, QCNE, US); along road of pine-plantation, 00°37'S, 78°27'W, 3435 m, 10 May 1984, S Laegaard 52107 (AAU, QCA, QCNE); 00°38'S, 78°33'W, 3400 m, S. Laegaard 69261 (AAU); 00°40'S, 78°30'W, 3900 m,Muhoz 207 (QCA); quebrada de Agualongo, 3400 m, 2 Feb 1992, Montesdeoca 280, 282, 284, 295, 339, Festuca in South American Paramos 348, 585 (AAU); 00°40'S, 78°30'W, 4550 m, Arguello 348 (QCA); 4300 m, Arguello 354, 357 (QCA); T. de Vries s.n. (AAU); Hacienda Pauzacha S of Volcan Cotopaxi, 00°44'S, 78°29'W, 3650 m, 2 Dec 1985, S. Laegaard 55734 (AAU, QCA); Cotopaxi, 3550 m, E. Asplund 6370 (AAU, F, MO, S, US); 3600 m, E. Asplund 6349 (AAU, F, MO, S, US); 3600 m, E. Asplund 6348 (AAU, F, S); road Zumbahua-Pujili, km 33, 00°53'S, 78°48'W, 3850-3900 m, 4 Apr 1992, S. Laegaard 102103 (AAU, QCA); 00°53'S, 78°48'W, 38503900 m, S. Laegaard 102100 (AAU); Angamarca road, km 5 from junction to road Latacunga-La Mana, 0POO'S, 78°55'W, 4000^1100 m, S. Laegaard 102134B (AAU); between Limpio Punja y Ruminahui, 00°38'S, 78°28'W, 3900^1200 m, S. Laegaard 55748 (AAU); E of Pilalo, km 19, 3600 m, 15 Apr 1990, P.M. Peterson 8759, C.R. Annable & M.E. Poston (K, MO, QCA, QCNE, US); km 21.5, 3820 m, P.M. Peterson 8770, C.R. Annable & M.E. Poston (MO, QCA, QCNE, US); SW of El Chaupi, km 6.6 on road to base of Illinizas, 3800 m, 1 May 1990, P.M. Peterson 8951 & C.R. Annable (K, MO, QCA, QCNE, US); Canton Latacunga-VolcanRutzalagua, 00°57'S, 78°33'W, 2900-3500 m, 14 Feb 1994, Ceron et al. 25697 (QAP); Paramo de Apagua entre Zumbagua y Pilalo, 4200 m, Barclay & Juajibioy 8085 (MO, US); Salcedo-Napo road, 00°55'S, 78°28'W, 3800-3850 m, J. Luteyn 13452 (MO, NY, QCA, QCNE, US); Pilalo-Latacunga road, 00°57'S, 78°58'W, 3400 m, Holm-Nielsen 1485 (AAU, NY, US); road Quevedo-Latacunga, Zumbagua, 3700 m, Hading et al. 8924 (GB); Mulatos paramo, lagunas, 13000 ft, Prescott 656 (NY); Panamerican Hwy, km 6 A Lasso, quebrada vegetation ENE Pastocalle, 3400 m, Sparre 15828 (S); Mun. Chaupi, NE slope of volcan Illiniza Norte, 00°3 8' S, 78°43'W, 4000^1050 m, 12 Oct 2000, D. Stancik 4026, 4027, 4030, 4032 (PRC, QCA); Mun. Lasso, Parque Nacional Cotopaxi, NE side of volcano, 00°37.8'S, 78°24.26'W, 3850 m, 21 Jun 1999, D. Stancik 3119, 3146, 3147, 3149, 3150, 3151, 3152, 3153, 3155, 3884, 4026 (PRC, QCA); around Museum, 3500 m, D. Stancik 3158 (PRC, QCA); Parque Nacional Cotopaxi entrance, 00°39'S, 78°31'W, 3530 m, 28 Sep 2000, D. Stancik 3888, 3889 (PRC, QCA); SE slope of Cotopaxi, 00°37'S, 78°24.5'W, 3800-3900 m, 28 Sep 2000, D. Stancik 3882 (PRC, QCA). Imbabura: Paramo Mariano Acosta, road Yahuarcocha-Mariano Acosta, km 20, 00°20'S, 78°00'W, 3650-3750 m, 9 Feb 1992, 133 S. Laegaard 101181, 101164B (AAU, QCA); Laguna Grande de Mojanda, 00°08'S, 78°17'W, 3850 m, S. Laegaard 55635 (AAU); Volcan Cayambe, S side, 78°10'W, 00°15'N, 4300 m, Sklenaf & Kosteckovd 1881 (AAU); vicinity of Mojanda, 15 km N of Malchinguin, S of Otavalo, 4000^1500 m, 11 Oct 1974, Gentry 12685 (MO); Volcan Cotacachi, 00°22'N, 78°20'W, 42004300 m, S. Laegaard 54506 (AAU); Laguna Cuicocha, 3100-3300 m, Penafiel et al. 527, 667, 720 (MO, QCNE); 3100-3400 m, Penafiel et al. 402 (MO, QAP, QCNE); 3300-3350 m, Penafiel 1067 (QAP); Proantag, estrivaciones occidental de la Cordillera Oriental, 3500-3800 m, Acosta-Solis 19196 (US); Hacienda Yura Cruz, 10 km N of Ibarra, 00°22N, 78°05'W, 3700-3800 m, HolmNielsen 6462 (AAU, NY, MO, S); quebrada Rumipamba, 3700-4000 m, Ceron 29480 (QAP); Cayambe-Laguna San Marcos, 11200 ft, Cazalet & Pennington 5434 (K, NY, US); Cerro Imbabura, faldas occidentales, 2800-4000 m, Acosta-Solis 17670 (US); Mun. Urcuqui, road to Cerro Yanaurcu, 00°28'13"N, 78°18'45"W, 4100 m, 15 Oct 2000, D. Stancik 4089, 4101 (PRC, QCA); Mun. Cayambe, volcan Cayambe, 00°00'4"N, 78°01'40.4"W, 4200 m, 20 Oct 2000, D. Stancik 4163, 4164 (PRC, QCA); Mun. Otavalo, road from Otavalo to Laguna Mojanda, 00°07'57"N, 78°16'27"W,3800m, 19 Oct 2000,/). Stancik 4115 (PRC, QCA); Mun. Urcuqui, road to Cerro Yanaurcu, 00°28N, 78°17.5'W, 3900 m, 15 Oct 2000, D. Stancik 4096, 4164 (PRC, QCA). Loja: Road to Fierra Urcu, ca. 10 km from main road Loa-Saraguro, 03°33'S, 79°15'W, S Laegaard et al. 18868 (AAU, LOJA, QCA, QCNE); 03°43'S, 79°19'W, 3650 m, S. Laegaard & Sklenaf 20282A (AAU, LOJA); 03°'S, 79°19'W, 3520 m, Sklenaf & S. Laegaard 7092 (AAU); Cerro de Arcos, W of road Manu-Zaruma, 03°34'S, 79°28'W, 35003600 m, S. Laegaard et al. 20613 (AAU, LOJA); Mun. Saraguro, road to Fierra Urcu, 03°42'40"S, 79°18'12"W, 3400-3450 m, 24 Aug 2000, D. Stancik 3769, 3770, 3772, 3773, 3774 (PRC, QCA). Morona-Santiago: Hda. Huargualla-Hda. San Eduardo, way to Parque Nacional Sangay, 02°0.25'S, 78°27'W, 3700 m, 30 Jul 1999, D. Stancik 3326, 5527 (PRC, QCA); Parque Nacional Sangay, Plazabamba, 02°0.64'S, 78°26'W, 3600m,20 Jul 1999,/). Stancik3337(PRC, QCA); confluence of Quebrada Plazabamba Chico with Q. Plazanbamba, 02°0.48'S, 78°25.4'W, 3000 m, 21 Jul 1999, D. Stancik 3344 (PRC, QCA); ridge 134 Festuca in South American Paramos above Q. Plazabamba Chico, 3600 m, 20 Jul 1999, D. Stancik 3351 (PRC, QCA); Plaza Culebrillas, 01°58'S, 78°25'W, swampy plate with grassy vegetation and shrubby patches, 3500-3600 m, 22 Jul 1999, D. Stancik 3355 (PRC, QCA); Plaza Culebrillas, 01°58'S, 78°25'W, 3500-3600 m, 22 Jul 1999,Z>. Stancik 3353, 3354, 3360, 3363, 3367 (PRC, QCA). Napo: Road Salcedo-Napo, ca. 6 km NE of km 45, 3900 m, S. Laegaard 53373 (AAU, QCA, QCNE); Paramo de Miranda, 00°34'N, 77°39'W, 3700-3900 m, 23 May 1985, S. Laegaard 54409 (AAU, QCA, QCNE); 3900-4100 m, S. Laegaard 54416 (AAU, QCA, QCNE); Cerro Antisana, SW slope, 4200 m, Black 32 (AAU); below Laguna Micacocha, paramo de Loma Gorda, 3850 m, Holm-Nielsen 20789 (AAU); W side of Volcan Puntos, 00°12'S, 78°10'W, 4150-4200 m, S. Laegaard 54731 (AAU); Los Llanganati, entre Ainchilibi y Rio Portrero al E de Romo Paramo, 2500-3620 m, Barclay & Juajibioy 9160 (COL, US); 3600-3700 m, Barclay & Juajibioy 9142 (COL, MO, US). Pichincha: Volcan Pichincha, via occidental, 2800-3830 m, Barford & BlicherMathiesen 41546 (AAU, QCA, QCNE); 35005000 m, Gentry 12378 (MO, QCA); Guagua Pichincha, 4000 m, Harlington 4552 (S); 35004500 m, Ceron 28250 (QAP); N slope, ca. 5 km WSW of Cotocollao, 00°08'S, 78°33'W, Sparre 13709 (S); Sodiro s.n. (QPLS); Pichincha-Cerro Ventanillas, 00°09'S, 78°32'W, 3850 m, S. Laegaard 51060 (AAU, QCA, QCNE); Lloa-Guagua Pichincha, km 10, 4170 m, S. Laegaard et al. 102733 (AAU, QCA, QCNE); Sodiro s.n. (QPLS); Jameson 70 (K); Hacienda Montecielo on S slope, 3400-3800 m, Sparre 17408 (S); Jameson s.n. (K); 3400 m, E. Asplund 6134 (S); 8500 ft, Spruce 5509 (K, P, US, W); Sodiro s.n. (US); 3700 m, Balslev 23600 (AAU); N slope, ca. 5 km WSW Cotocollao, 3600-3750 m, Sparre 13709 (AAU); Volcan Pasachoa, 00°27'S, 78°30'W, 3700-3900 m, S Laegaard 55279 (AAU, COL, F, GB, K, LOJA, MO, NY, QCA, QCNE, VEN, W); 3700-3900 m, S. Laegaard 55279 (COL, MO); 00°27'S, 78°30'W, 3900 m, S. Laegaard 55276 (AAU, QCA); 00°27'S, 78°30'W, 3600 m, S. Laegaard 55286 (AAU); 3000-3500 m, Ceron & Alarcon 12260 (MO, PRC, QAP, QCNE); 3500^1300 m, Ceron & Alarcon 4805 (QCNE); 00°27'S, 78°28'W, 2850-3900 m, Paredes 41 (QCA); 3800 m, Penland & Summers 969 (F); Paramo Papallacta, 3900 m, Maf 57 (AAU); Guamani, 01°19'S, 78°12'W, 3600-3800 m, E. Asplund 8718 (QCA, S); 3200 m, Mille s.n. (QPLS); Mille 284 (US); E. Asplund 9634 (S); E. Asplund 8724 (MO, NY, QCA, S, US); 3900 m, E. Asplund 17176 (S); 4000 m, Vargas et al. 2083 (AAU, MO, QCNE); Ramsay 426 (K); 4200 m, Quintana 2 (QCA); 00°15'S, 78°12'W, 3800 m, 7 Oct 1987, Ramsay & Smith 742 (K, QCA, QCNE); 3800 m, Harling et al. 10477 (GB); Andibus Quitensis, Jameson s.n. (K, S); vicinity of Quito, 3200 m, E. Asplund 6140 (S); Quito-Andes, Jameson 91 (K, W); 3500 m, E. Asplund 6138 (F, S); Volcan Antisana, falda W, 4200 m, Ehrenburg 191 (QCA); 4150 m, Grubb et al. 656 (NY, K); W side, 00°28 'S, 78°12'W, 4200 m, Balslev et al. 4070 (QCA); Hacianda de Antisana, Hartweg 1455 (K, NY, US); between campamento IMAP and Laguna Micacocha, 00°33'S, 78°12'W, 3850-3950 m, S. Laegaard 101577 (AAU); between Mt. Antisana and Mt. Sincholagua, E. Asplund 8647 (NY, QCA, S); E. Asplund 8645 (S); E. Asplund 8644 (QCA, S); road from Otavalo to Laguna Mojanda, 3810m, J.F. Smith 1997 (AAU, F, MO, NY, QCNE, US); J.F. Smith 1999 (NY, QCA, QCNE); D. Stancik 4103 (PRC); 00°08'S, 78°16'W, 3725-3750 m, S. Laegaard 52373 (AAU, QCA); Laguna Negra, 00°08'S, 78°16'W, 3800 m, S Laegaard 54351 (AAU, QCA, QCNE); Illiniza Sur, E slope, 00°40'S, 48°42'W, 4200 m, Sklendf & Kosteckovd 14-9 (US); Illiniza Norte, 4000-4100 m, Sparre 15781 (S); Mount Atacazo, 4700 m, Harteman 67 (US); 00°02'N, 78°37'W, 4250 m, S. Laegaard 53871 (AAU, QCA, QCNE); 00°20'S, 78°38'W, 37503800 m, S Laegaard 55672 (AAU, QCA), SW slope, km 19 from San Juan, 00°21'S, 78°39'W, 2900 m, Holm-Nielsen 25163 (AAU); 00°20'S, 78°36'W, 3600 m, Espinosa 81 (QCA); road Chillogalo-San Juan, 00°18'S, 78°39'W, 31003260 m, 8 Sep 1945, Jaramillo & Zak 623 (AAU, MO, PRC, QCA); Sincholagua, E. Asplund 8639 (QCA, S); Pululahua-San Bartolo al E de Moraspungo, 00°01'S, 78°29'W, 3050 m, 13 Dec 1990, Ceron & Montesdoca 12010 (QAP); Mont Ungui, 3400 m, Firmin 137 (F, US, W); El Corazon, 11000 ft, Prescott 834, 839, 848 (NY); 4100 m, E. Asplund 17522 (S); E side of Corazon, 1314,000 ft, Feb 1880, Whymper 1322 (K); Pogonales de Pichincha, 13 May 1930, Benoist2515 (P); Mun. Pifo, Paramo de Guamani, 00°19'S, 78°15'W, 3700 m, 19 Jun 1999, D. Stancik 3013 (PRC, QCA, W); Mun. Amaguana, E slope of volcan Pasachoa, 00°27'56"S, 78°28'40"W, 4150 m, 14 Sep 2000, D. Stancik 3671, 3677, 3687, 3688, 3690, 3691, 3692, 3704 (PRC, QCA). Sucumbios: SW de Festuca in South American Paramos Playon de San Francisco, Paramo Mirador, 34003600 m, 15 May 1990, P.M. Peterson 9155, E.J. Judziewicz & R.M. King (MO, QCA, QCNE, US). Tungurahua: Cordillera de Llanganatis, Lake Yanacocha, 3600 m, E. Asplund 9967 (QCA); El Tambo near Lake Yanacocha, 3650 m, E. Asplund 9964 (S); Parque Nacional Llanganatis, via Salcedo-Tena; 3600 m, 17 Apr 1998, Vargas etal. 2583 (MO, QCNE); Lake Aucacocha, 3700 m, Aug 1969, Edwards 70 (K); Las Tolas-5 km SE of LagunaPisayambo, 01°07'S, 78°21 'W, 3700 m, 12 Jan 1999, S Laegaard & Grignon 19395 (AAU, LOJA, QCA); road Ambato-Guaranda, 01°25'S, 78°51 'W, 4070 m, Brandbyge 42133 (AAU, QCA); Comunidad Rumipata, 00°22 'S, 78°55' W, 4000 m, Brandbyge 42586 (MO, AAU); Cerro Sangay, 4000 m, Stubel 257 (S); Mun. Pillaro, Las Llanganatis, around Aucacocha lagoon, 01°9'S, 78°20'W, 3800 m, 28 Sep 2000, D. Stancik 3901, 3902 (PRC, QCA); Las llanganatis-Pisayambo, 01°07'S, 78°20'W, 3600 m, 28 Sep 2000, D. Stancik 3900 (PRC, QCA). PERU. Amazonas: Prov. Chachapoyas, encima de Leimebamba, pajonal, 3400-3500 m, 17 Mar 1964, R- Ferreyra 15518 (US); upper slope and summit of Cerro Campanario, 3600-3900 m, 3 Aug 1962, J. Wurdack 1571 (US). Ancash: Prov. Huaylas, Huascaran Parque Nacional, quebrada Alpamayo at foot of snowfree peak above Lago Jancarirish, 08°53'S, 77°41 'W, 4350^1500 m, 3 Sep 1985, Smith et al. 9783 (MO); Prov. Recuay, Huascaran Parque Nacional, Rio Pachacoto drainage between mineral springs and Pumashimi, 09°53 'S, 77° 17'W, 42004300 m, 16 Mar 1986, Smith et al. 11799 (MO); Prov. Yungay, Huascaran Parque Nacional, Llanganuco sector, quebrada Demanda, trail to Chacraraju base camp and Brogue glacier, 09°02'S, 77°36'W, 4100-4400 m, 4 Dec 1985, Smith et al. 10253 (MO); Huascaran Parque Nacional, Llanganuco sector, quebrada Ancosh, 09°03'S, 77°35'W, 4500 m, 4 Feb 1985, Smith et al. 10229 (MO). Cajamarca: Prov. Cajamarca, dist. Cajamarca, entre Cajamarca y Cumbe Mayo, km 14, Al arboretum Cumbe Mayo de Cicafor, 3400 m, 18 Apr 1981, Vega et al. 2464 (F); Cajamarca-Bambamarca road, in puna, 07°00'S, 78°33'W, 3800 m, 17 Feb 1983, Smith 3482 (MO); Bajando el Paso El Gavilan hacia San Juan, el E de la carretera Cajamarca-Pacasmayo, 3000 m, 18 May 1986, Vega 4043 (F); a la altura del Paso El GavilanGavilan, 3200 m, 18 Apr 1976, Vega 1385 (F); Jalca de Kumullca, entre La Encanada y Celendin, 3600 m, 135 27 May 1977, Vega et al. 2011 (F); Sorochuco, Carretera Michiquillay-El Punre, 07°00'S, 78°18'W, 3570 m, 9 Aug 2001, Vega et al. 10904 (F); 9 Sep 2001, Vega etal. 10906 (F); humid jalca vegetation with tussock grasses, 07°02 'S, 78° 15 'W, 26 Aug 2004, Sklendf & Zapata Cruz 8630 (PRC); Cumbe Mayo, 21 km al W de Cajamarca, ladera con arbustos dispersos, 3100 m, 4 Nov 1977, Vega 1969 (F); Coymolache ruta Cajamarca-Hualgalloc, 3850 m, 7 Jan 1977, Vega et al. 2052 (F); Pampa Larga, al N de la explotacion minera Yanacocha, ladera graminosa de jalca, 3900 m, 14 May 1994, Vega 7136 (F); Prov. Hualgayoc, desvio de la carretera Coymolache-Chugur, jalca graminosa, 3700 m, 28 Apr 1994, Vega et al. 7062 (F); Prov. Hualgayoc, Coymolache, 06°46'S, 78°38'W, 4000 m, 15 Jun 2001, Vega et al. 10795 (F, MO); Prov. Hualgayoc, El Tingo, desvio a Chugur, 3350 m, 22 Jul 1986, Vega 4145 (F). Cuzco. Paso de Tres Cruces, Cerro de Cusilluyoc, 3800-3900 m, 5 Mar 1925, Pennell 13845 (NY); Sacsahuaman, rocky stream-bank, 3500-3600 m, 24 Apr 1925, Pennell 13552 (NY); Prov. Espinar, Hda Cuyo, 4200^1500 m, 24 Mar 1956, Vargas 111 76 (US). La Libertad. Prov. Bolivar, ascenso ao Nevado de Cajamarquilla, 07°08'S, 77°42'W, 3000 m, Vega 11181 (F, MO); entre el desvio a Uchumarca y Santa Luisa, 07°04' S, 77°49'W, 3700 m, Vega 11190 (F, MO); Prov. Otuzco, Trujillo-Huamachuco road, 10-15 km before Shorey, 07°59'S, 78°22'W, 3300 m, 13 Feb 1983, Smith et al. 3278 (MO); Prov. Pataz, grassland in Manachaqui valley, ca. 10 km NE of Pataz, 07°40'S, 77°30'W, 3600 m, 22 Mar 1988, B. Leon & Young 1679 (MO); Manachaqui valley, ca. 10 km NE of Pataz, 77°30'W, 07°40'S, 3600 m, 28 Feb 1988, B. Leon & Young 1101 (MO); Prov. Santiago de Chuco, al oeste del cementerio de Quiruvilca, 3870 m, 21 May 2001, Leiva & Leiva 2525 (MO); Paso de Alaska. Carretera a Tayabamba, 3900 m, 24 Jun 1974, Lopez & Sagdsteguii 8180 (MO, US). San Martin. Prov. Huallaga, distr. Saposoa, entre la Artesa y Rangra Pata, 3600-3800 m, 23 Aug 2001, Quipuscoa & Vilchez 2611 (F, MO); Prov. Mariscal Caceres, Pampa de Cuy, NW sector Rio Abiseo Parque Nacional, 07°40'S, 77°30'W, 3550 m, 3 Jun 1988, B. Leon & Young 1364 (MO); Chochos valley, NW corner of Rio Abiseo Parque Nacional, 3425 m, 31 May 1986, Young & Watson 3484 (MO). Note. Ocassionally there are proliferating plants with clearly transformed spikelets and sexual organs. Panicles are compact with short deformed branches; 136 Festuca in South American Paramos spikelets are unisexual, transformed into vegetative shoots, only exceptionally with some spikelets carrying rudiments of sexual organs; and some florets are without anthers and ovaries. These plants can be distinguished from F. asplundii by having narrower leaf blades (0.4•0.8 versus 1-1.4 mm wide) and viviparous panicles that are abnormally developed (versus normally developed in F. asplundii). Viviparous specimens examined. COLOMBIA. Nariflo: Mun. Tuquerres, Volcan Azufral, road from vereda San Roque Alto to Laguna Verde, km 6, 3850 m, 9 Mar 1999, D. Stancik 2790 (COL, PRC, PSO). ECUADOR. Morona-Santiago: Parque Nacional Sangay, Plazabamba, 02°00.64'S, 78; 26'W, 3600 m, 19 Jul 1999, D. Stancik 3335, 3336 (PRC, QCA). Pastaza: E of pass Alao-Huamboya, 01°48'S, 78°25'W, 3700-3900 m, S. Laegaard 55452 (AAU, PRC, QCA). Tungurahua: Llanganatis, Las Tolas, 5 km SE of Laguna Pisayambo, 01°07'S, 78°21'W, 3700 m, 12 Jan 1999, S. Laegaard 19395 (AAU, LOJA, QCA); S of Laguna Verde at Cerro Hermoso, 01°14'S, 78°18'W, 3850 m, 11 Nov 1980, Holm-Nielsen & Jaramillo 28415 (AAU, QCA). 53. Festuca sumapana Stancik, Darwiniana 41(1-4): 144, f. 71-p. 2003. (Figs. 67, 68). TYPE: Colombia. Meta, Paramo de Sumapaz, Cerro Nevado del Sumapaz, 4130 m, 16 Jan 1973, Cleef7930 (holotype: COL!; isotypes: U!,US!). Tussocked perennials with intravaginal innovations. Culms 15-20 cm tall, erect, glabrous; nodes 1, basal and 1 leaf. Leaf sheaths membranous, grayishstramineous, glabrous; auricles absent; ligules 0.3-0.5 mm long, membranous, apex two-lobate; blades 8-12 cm long, 0.5-0.7 mm wide, involute, abaxially scabrous, green, apex obtuse. Panicles 5-7 x 0.4•0.6 cm, lanceolate, contracted; branches scabrous. Spikelets 9.5-11 mm long, lanceolate, florets 3 or 4; rachilla glabrous; glumes 4•6 mm long, lanceolate, keeled, membranous, purplishgreen, scabrous dorsally, apex acute; lower glumes 4•5 mm long, 1-nerved; upper glumes 5.5-6 mm long, 3-nerved; lemmas 6.5-7 mm long, lanceolate, 5-nerved, keeled, membranous, purplish-green, papillose, awned, the awn 0.5-1.5 mm long; callus glabrous; paleas as long as the lemma, lanceolate, membranous, margins scabrous; lodicules lanceolate; anthers 0.6-0.8 mm long; ovary apex glabrous. Caryopses not seen. Leaf blade anatomy.•Cross-sections usually with 5 vascular bundles, 3 ribs above; sclerenchyma under abaxial epidermis discontinuous, adaxially absent; adaxial epidermis with scattered hairs, the hairs ca. 0.07 mm long. Observations.•Festuca sumapana is morphologically similar to F glumosa but differs in having shorter ligules (0.3-0.5 versus 0.8-1.2 mm), narrower leaf blades (0.5-0.7 versus 0.8-1.4 mm), and shorter lower glumes (4-5 versus 4.5-6 mm). Distribution and habitat.•Festuca sumapana is endemic to Colombia where it is known only from the type locality, Paramo de Sumapaz (Cundinamarca). It occurs at an altitude about 4000 m in the superparamo zone. 54. Festuca tolucensis Kunth, Nov. Gen. Sp. (quarto ed.) 1:153.1816. (Figs. 67,69,97A-C). TYPE: Cresit in montosis, scopulosis, apricis regni Mexicani, inter Islahuaca et Toluca, 1380 hexap., Sep, Humboldt& Bonplands.n. (holotype: B!; isotypes: B!, BM, P!). Festuca aequipaleata E. Fourn., Biol. Cent.-Amer, Bot. 3(20): 581. 1885. Festuca aequipaleata E. Fourn., Mexic. PI. 2: 125. 1886, isonym. TYPE: Mexico. Orizaba, 14,000 ft, Liebmann 6108 (lectotype: C!, designated by Alexeev, Novosti Sist. Vyss. Rast. 21: 47. 1984; isolectotypes: C!, K!). Festuca liebmannii E. Fourn., Mexic. PI. 2: 124. 1886. TYPE: Mexico. Liebmann 517 (holotype: P; isotype: US!). Festuca multiculmis Steud., Syn. PI. Glumac. 1: 310. 1854. TYPE: Mexico. Mt. Toluca, Heller 306 (holotype: P!). Tussocked perennials with intravaginal innovations. Culms 60-80 cm tall, erect, finely scabrous; nodes 1, basal. Leaf sheaths membranous-coriaceous, stramineous to grayish, glabrous; auricles absent; ligules 1.8-3.5 mm long, membranous, apex acute; blades 20-30 cm long, 0.5-0.7 mm wide, involute, green to olive-green, abaxially scabrous, apex acute. Panicles 20-25 x 0.5-1.5 cm, lanceolate, contracted, branches finely scabrous. Spikelets 7•12(•15) mm long, lanceolate sometimes oblong-lanceolate, florets 3 or 4(-6); rachilla densely hairy; glumes 6-9.5 mm long, lanceolate, coriaceous, green, markedly scabrous, apex acute; lower glumes (2-)6-8.5 mm long, 1-nerved; upper glumes (4•)6.5-9.5 mm long, 3-nerved; lemmas 6-9.5(-10.5) mm long, lanceolate, 5-nerved, coriaceous, green, scabrous on all surfaces, Festuca in South American Paramos 137 2 mm Figure 68. Festuca sumapana. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Clee/7930 (COL). 138 Festuca in South American Paramos 1 mm Figure 69. Festuca tolucensis subsp. tolucensis. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 4279 (PRC). Festuca in South American Paramos awned, the awn 1-1.5 mm long; callus glabrous; paleas almost as long as the lemma, lanceolate, membranous, scabrous, short-hairy; anthers 2.5-3 mm long; ovary apex glabrous. Caryopses lanceolate; hilum 5/6 as long as the grain, linear. Leafblade anatomy.•Cross-sections typically with 5-9 vascular bundles and 3-5 ribs above; sclerenchyma under abaxial epidermis continuous, extending to somevascular bundles, adaxial sclerenchyma present in 3-5 ribs, isolated; abaxial epidermis with a dense covering of prickles; adaxial epidermis hairy, the hairs 0.03-0.05 mm long. Observations.•Festuca tolucensis is morphologically similar to F. subulifolia and both species 139 form tussocks of fine-leaved plants. Besides differeing in leaf cross-section, F. tolucensis has spikelets 7•12(•15) mm long (versus 6.5-9 in F subulifolia), lower glumes 6.5-9.5 (versus 3.5-5.5) mm long, and upper glumes 6.5-9.5 (versus 4-6) mm long. Distribution and habitat.•Festuca tolucensis has a wide distribution and ranges from Mexico to Venezuela. In Colombia, Festuca tolucensis is only known from the Serrania de Perija. We recognize three subspecies as differentiated in the tabular comparison and key below. It is a dominant species found in dry rocky slopes or swampy margins of lagoons in grass paramos between 3000-4000 m. F. tolucensis subsp. Distribution Glume length (mm) Lemma Lower Upper Length (mm) Spikelets No. florets Length (mm) Vascular bundles No. Girders tolucensis perijae culata MEX-VEN COL, VEN VEN (4-)5-6 5.5-6.5 2-2.5 4-5 2 or 3 4 or 5 7-9 5 5 (5-)6-8 6.5-7.5 4-5 6.5-7.5 6-7 6-7 7-10 9-10 10-11 l-3(-5) 0 1 KEY TO THE SUBSPECIES OF FESTUCA TOLUCENSIS la. Lower glumes 2-2.5 mm long; upper glumes 4-5 mm long; spikelets 10-11 mm long 54c. F. tolucensis subsp. culata lb. Lower glumes (4-)5-6.5 mm long; upper glumes (5-)6-8 mm long; spikelets 7-10 mm long.... 2 2a. Small tussocks (less than 40 cm tall), leaves with 5 vascular bundles, spikelets with 2 or 3 florets; 9-10 mm long 54b. F. tolucensis subsp. perijae 2b. Tussocks more than 40 cm tall, leaves with 7-9 vascular bundles, spikelets with 4-5 florets; 7-10 mm long 54a. F. tolucensis subsp. tolucensis 54a. Festuca tolucensis subsp. tolucensis (Fig. 69). Additional specimens examined. COSTA RICA. San Jose: Iraza, 1300 m, 14 Jun 1932, Storek 2889 (MO); Canton de Perez Zeledon, Parque Nacional Chirripo, Cordillera de Talamanca, Rio Talari, cerca del cruce al Cerro Ventisqueros, 3600-3450 m, 27 Jan 1996, Morales 5194 (MO); Canton de Perez Zeledon, Parque Nacional Chirripo, cuenca Terraba-Sierpe Chirripo, base Crestones, 09°27.5 'N, 83°30.5'W, 3460 m, 12 Jul 1996, Alfaro 1029 (MO); Canton de Perez Zeledon, Parque Nacional Chirripo, cuenca Terraba-Sierpe, Sandero a Valle Los Leones, 09°26'N, 83°30.5'W, 3100-3400 m, 18 Jul 1998, Alfaro 1771 (MO). MEXICO. D.F.: Ajusco, Sep 1928, Lyonnet 260 (NY); Lomas, Oct 1932, Lyonnet 2973 (MO). Volcan Toluca, 4150 m, 18 Oct 1953, Sohns & Matuda 991 (NY); W side in volcanic ash, competing with Sedum minimum, 3940 m, 9 Nov 1955, J. Clausen 34 (NY); crater, above timberline, dwarf alpine plants, 14000 ft, 17 Aug 1972, Dziekanowski & Bolingbroke 1911 (MO, NY); 1846, Heller 62, 306 (P, W); 4150 m, 18 Oct 1953, Sohn & Matuda 991 (P). Iztaccihuatl, side of Mt. at La Joya, 3990 m, 7 Jun 1960, J. Beaman 3482 (NY); Monte de Rio Frio, km 49, road from Mexico City to Puebla, 4000 m, 31 Jul 1929, Mexia 2684 (NY); Mt. Popocatepetl, 3400 m, 8 Jun 1910, A.S. Hitchcock 496 (C, MO, NY, W); Tlaxcala vertiente N del Volcano La Malinche, zacatonal de Festuca y Calamagrostis, 3920 m, 16 Sep 1986, Gonzalez et al. 206 (F, MO); Mun. de Amecameca, bosque de 140 Festuca in South American Paramos pino, 3900 m, 30Nov 1980, Juan &Alva 101 (MO). Chiapas: Summit of Volcano Tacana, Mun. Union Juarez, scattered dwarfed Pinus sp., 2200 m, 30 Jul 1972, D. Breedlove 26723 (NY); Volcan Tacana, near summit sandy, gravelly soil above timberline, 4090 m, 8 Dec 1959, J. Beaman 3216 (NY); Mt. Tacana, 2000-4038 m, VIII 1938, Matuda 2337, 2360 (NY). Jalisco: Nevado de Colima, side of mountain below Puerto de Colima, sandy soil in sparsely vegetated meadow, 4000 m, 26 Aug 1958, J. Beaman 2363 (NY); S of Ciudad Guzman (Zapotlan), near summit, in rockslide, gravel, around rocks, 4000 m, 7 Feb 1956, Gregory & Eiten 301 (MO, NY, P); Mt. Nevada, 14300 ft, 23-24 Sep \9\0,A.S. Hitchcock 7165 (NY). Michoacan: Tancitaro region, Mt. Tancitaro, 10000-11000 ft, 25 Jul 1941, Leavenworth & Hoogstraal 1218 (MO, NY). Puebla: Arroyo Paso Buey, NW side of volcano Pico de Orizaba and 7 km NW of the summit, 0.5 km SE of Miguel Hidalgo, 97°18'30"W, 19°04'30"N, 3300 m, 9 Jul 1986, M. Nee & Soule 33022 (MO, NY); Faldas del Pico Orizaba, 3300 m, 17 Oct 1971, Hernandez et al. 1336 (F, MO). Veracruz: Mun. Perote, km 1-2 above Escobillo on the NW slopes of Cofre de Perote 3300 m, 21 Jan 1984, Taylor et al. 170 (F, MO, NY); Pico Orizaba, NE side of Mt., 3980 m, 16 Aug 1958, J. Beaman 2284 (NY); Mt. Orizaba, sandy plains, 14000 ft, 7 Aug 1891, Seaton 228 (NY); 18 km de Perote camino a la cima del Cofre, 3790 m, 10 Feb 1984, Gonzalez et al. 137 (MO); Pico de Orizaba, 14000 ft, Sep 1941,Liebmann2886, 3030, 6117(C); Wawra 947 (W); Ross 1276 (W). VENEZUELA. Merida: Sierra Nevada-Laguna La Coromoto, 3300-3400 m, 18 Feb 1966, Schulz & Rodriguez 317 (US); 13000 ft, Feb 1846, Funck & Schlim 1132 (P); 4000-4400 m, 24 Sep 1952, Humbert 26353 (COL, MER, P, US); 4000-4400 m, Humbert 26806 (MER); Humbert 26506 (P, US); pastizal, 3700 m, 25 Sep 1970, Castelano & Monasterio 20 (VEN); Laguna La Coromoto, Plantaginetum, 3200 m, 24 Feb 1955, Vareschii 3853 (VEN); 3400 m, Oct 1956, Aristeguieta 2602 (VEN, US); 3300 m, 3 Jul 1987, Briceno & Adamo 1988 (MERF); Valle de Mucubaji, abajo del Pico Mucunuque, 08°46'35"N, 70°48 '57"W, 3730 m, 3 Sep 1998, Berg 98-29-16 (K); Laguna Mucubaji, swamps around the lagoon, 8°47.5'N, 70°49'W, 3600 m, 6 Nov 2000, D. Stancik 4172 (CAR, COL, PRC); 3570 m, 23 Aug 1980, Briceno & Adamo 184 (Herbarium Briceno, MERF, MO, PRC, US); camino hacia Mucunuque, 4000-4300 m,4Nov l992,Meier et al. 3026 (VEN); Laguna de Mucubaji, 3600 m, Dec \952, Aristeguieta 964 (US); Paramo de Mucub- aji, 3500 m, 23 Aug 1980, Briceno & Adamo 187 (MO); S side of Hwy 7, 5 km E of Laguna Victoria, 3000 m, 22 Aug 1972, Hanselmann & Loleveless 296 (MO); Laguna de Mucubaji, 8 Nov 1952, Aristeguieta s.n. (VEN); 3900 m, 9 Sep 1958, Vareschi 7021 (VEN); 8 Nov 1952, Aristeguieta 947 (VEN). Mun. Tabay, Laguna La Coromoto, 8°35.5'N, 71°01'W, 3300 m, 7 Nov 2000, D. Stancik 4181 (CAR, COL, PRC); Laguna La Coromoto, 3300-3400 m, 15 Dec 1966, Schulz 317 (MER); Sierra St. Domingo, Pico de Mucunuqui, al S de Laguna Grande, 4400 m, 28 Nov 1959, Barclay & Juajibioy 9895 (COL); Paramo de Mucuchies, Pico de Aguila, 4418 m, 21 Nov 1959, Barclay & Juajibioy 9656 (COL, MO, US); Sierra del Norte, 6 Oct 1952, Humbert 26824 (COL, MER, P, US); Mun. Mucuchies, Paramo de Piedras Blancas, Lagua Negra, 8°49'N, 70°57'W, 4270 m, 4 Nov 2000, D. Stancik 4191 (CAR, COL, PRC, W); 4200 m, D. Stancik 4207 (CAR, COL, PRC, W); D. Stancik 4176 (AAU, CAR, COL, PRC); 8°48.5'N, 70°55.5'W, 3800 m, D. Stancik 4225 (CAR, COL, PRC); Laguna Tapada, 8°49'N, 70°5643'W, 4100 m, D. Stancik 4242 (AAU, CAR, COL, PRC); 4100 m, D. Stancik 4243 (CAR, COL, PRC, W); Laguna Negra, 9 Sep 1952, Aristeguieta 1003 (VEN); Laguna Los Patos, Laguna El Infernito, 3700 m, Aug 1956, Aristeguieta 2464 (VEN). Paramo de Piedras Blancas, 4400 m, 8 Dec 1979, Barreto 645 (MERC); 4000^1600 m, 9 May 1985, Briceno et al. 1146 (MERC); 4000^1600 m, 14 Nov 1981, Briceno et al. 440 (Herbarium Briceno, MERC); sector Las Cruces, 3700-3900 m, 8 Sep 2000, Briceno 3711 (Herbarium Briceno); Mun. La Culata, Paramo La Culata, Briceno & Adamo 1067 (MERF); 3300 m, 7 Jun 1984, Briceno & Adamo 949 (MERF); Mun. Mucuchies, Paramo de Misinta, NE de Mucuchies, 4000 m, 26 Oct 1984, Briceno & Adamo 1139 (MERF); Paramo Las Monsalves, selva de Polylepis, 3900 m, 10 Nov 1952, Vareschi 2232, 2134, 2244 (VEN); 3920 m, Vareschi 2248 (VEN); Pico Espejo-Timoncito, paramo, 4300^1700 m, 16 Dec 1952, Bernardi 275 (NY). Tachira: Mun. La Grita, Paramo La Negra, 08° 15 'N, 71 °53' W, 3200 m, 11 Nov 2000, D. Stancik 4280 (CAR, PRC); D. Stancik 4279 (CAR, COL, PRC); Sierra Nevada, camino a Pico Bolivar, cerca de Refugio Moya, 4100-4300 m, 10 Oct 1953, Littele 15712 (MER); Pico Bolivar y Espejo, 15-18 Dec 1959, Barclay & Juajibioy 10214 (US); Sierra Nevada, Loma Redonda, Paramo Media Luna, 4100 m, 3 Feb 1995, Berg 649 (MERC); Berg & Steinmetz 187 (Herbarium Briceno, MERC); Loma Redonda, 3940 m, Festuca in South American Paramos 7 Oct 1994, Briceno 2854 (Herbarium Briceno); Alto de La Cruz, 4040^1300 m, 11 Nov 1994, Briceno et al. 3077 (Herbarium Briceno); Mun. Rangel, Laguna La Ciega, 4200 m, 29 Sep 2000, Briceno & Molinillo 3766 (Herbarium Briceno); Mun. Rangel, Paramo El Banco, 4420 m, 21 Oct 1997, Briceno & Molinillo 3447 (Herbarium Briceno); Paramo de Misinta, NE de Mucuchies, 4000 m, 5 Jun 1987, Briceno & Adamo 1131 (Herbarium Briceno); Paramo El Toro, Los Arbolitos y Los Nevados, 3100 m, 12 Oct 1988, Briceno et al. 2330 (Herbarium Briceno); cresta Pico el Toro, 4200 m, 25 Nov 1994, Briceno 3119 (Herbarium Briceno, PRC); via El Leon, 3460-3580 m, 4 Nov 1994, Briceno 3111 (Herbarium Briceno, PRC); Valle de los Calderones, Paramo de los Pozones, 3350 m, 19 Oct 1995, Briceno 3357 (Herbarium Briceno); Laguna Verde-Laguna Suero, 3990-4190 m, 29 Mar 1994, Briceno et al. 2667 (Herbarium Briceno); Laguna El Espejo, 3800-3950 m, 9 Dec 1994, Briceno 3178 (Herbarium Briceno); Laguna de Los Anteojos, 3900 m, 2 Dec 1994, Briceno et al. 3158 (Herbarium Briceno); Laguna La Fria, 3460-3580 m, 4 Nov 1994, Briceno et al. 3054 (Herbarium Briceno, PRC). Trujillo: Paramo de la Cristalina, 2900 m, Oct 1910, Jahn 18 (US, VEN); 20 Oct 1910, Jahn 43 (VEN); Tuname-Guirigoy, 3200 m, Aug \95%, Aristeguieta& Medina 3531 (VEN); Guirigoy, hacia La Parida, 3400 m, Aug 1958, Aristeguieta & Medina 3558 (VEN); Dept. Escuque, Cabimbi, Teta de Niquitao-Laguna Negra, 3800-3900 m, 19 Aug 1988, Briceno s.n. (Herbarium Briceno). 54b. Festuca tolucensis subsp. perijae Stancik, Darwiniana 41(1-4): 146, f. lla-f. 2003. TYPE: Colombia. Magdalena, Sierra de Perija, E of Manaure, Cerro Avion, 3550-3450 m, 8 Nov 1959, J. Cuatrecasas & Castaneda 25136 (holotype: COL!; isotype: US!). Panicles slender and compact. Spikelets 9-10 mm long; lower glumes 5.5-6.5 mm; upper glumes 6.5-7.5 mm long; lemmas 6-7 mm long. Leafblade anatomy.•Abaxial epidermis continuous, not extending to the vascular bundles. Distribution and habitat.•Fesctuca tolucensis subsp. perijae is known only from the Serrania de Perija on the Colombian-Venezuelan border, and is found in paramos at an altitude between 3000-3500 m. Additional specimens examined. COLOMBIA. Magdalena: Sierra de Perija, plain between Cerro Venado and Cerro Avion, 3270-3350 m, J. Cuatreca- 141 sas & Castaneda 25133 (COL, US). VENEZUELA. Zulia: Sierra de Perija-Serrania de Valledupar, campamento "Monte Viruela", 10°25 '13"N, 72°52 '42"W, 3100 m, S.S Tillett 747-1122 (COL). 54c. Festuca tolucensis subsp. culata Stancik & P.M. Peterson, subsp. nov. TYPE: Venezuela. Merida, Mun. La Culata, Paramo La Culata, 3300 m, 8°45'42"N, 71°03'W, 3300 m, 12 Nov 2000, D. Stancik 4259 (holotype: PRC!; isotypes: CAR!, COL!). Haec subspecies a Festuca tolucensis subsp. tolucensis et F tolucensis subsp. perijae spiculis 10-11 (non 7-10) mm, glumis minoribus (inferioribus 2-2.5 non 4-6.5 mm, superioribus 4-5 non 6-7.5 mm) distinguitur. Spikelets 10-11 mm long; lower glumes 2-2.5 mm long; upper glumes 4-5 mm long. Leaf blade anatomy.•Abaxial epidermis continuous, extending just to a single vascular bundle. Distribution and habitat.•Festuca tolucensis subsp. culata is known only from Merida where the subspecies occurs in rocky paramos of La Culata and El Escorial, at an altutude of about 3000-3350 m. Additional specimens examined. VENEZUELA. Merida: Paramo El Escorial, 3100-3420 m, 10 Jul 1987, Briceno & Adamo 2052 (Herbarium Briceno, PRC). Mun. La Culata, Paramo La Culata, 3300 m, 8°45'42"N, 71°03'W, 3300 m, 12 Nov 2000, D. Stancik 4258 (CAR, COL, PRC). 55. Festuca turimiquirensis Stancik & P.M. Peterson, sp. nov. (Figs. 49, 70). TYPE: Venezuela. Anzoategui, Distr. Libertad, ridges and tops of Montanas Negras, along the Sucre and Anzoategui border, 20 km NE of Bergantin, NE of Buenos Aires, Serrania de Turimiquire, 10°04'30"N, 64° 11 'W, 2000-2350 m, bambooericaceous scrub and elfin cloud forest, 28 Nov 1981, G. Davidseetal. 19610(holotype: MO!, isotypes: NY!, VEN!). Haec species a F tolucensis glumis minoribus (inferibus 3•4 non 5-8.5 mm, superioribus 4-5.5 non 5-9.5 mm), aristis longioribus et paniculae majoribus (6-7 cm lat., non 0.5-1.5 cm), et sclerenchymate abaxiali non continui differ! Tussocked perennials with intra- and extravaginal innovations. Culms 70-80 cm tall, erect, 142 Festuca in South American Paramos 100 x Figure 70. Festuca turimiquirensis. A. Stylized growth form. B. Habit. C. Extravaginal shoot. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Caryopsis. J. Leaf blade cross-section. A-I, Davidse et al. 19610 (NY). Festuca in South American Paramos glabrous; nodes 3 or 4 near base. Leaf sheaths membranous, brown, glabrous, striate; ligules 1.5-4 mm long, apex acute; blades 40-50 cm long, 0.6-0.7 mm wide, conduplicate, abaxially glabrous, green. Panicles 15-17 x 6-7 cm, open, ovate; branches glabrous. Spikelets 10-11 mm long, ovate, florets 4; rachillas 1.2-1.4 mm long, hairy; glumes 3-5.5 mm long, membranous, upper 1/3 scabrous, apex acute; lower glumes 3•4 mm long, lanceolate, 1-nerved; upper glumes 4-5.5 mm long, lanceolate, 3-nerved; lemmas 7-8.5 mm long, lanceolate, 5-nerved, membranous, upper 1/2 scabrous, awned, the awn 0.7-2.5 mm long; paleas as long as the lemma, membranous, scabrous; anthers ca. 3 mm long; ovary apex sparsely hairy. Caryopses lanceolate; hilum 5/6 as long as the grain, linear. Leaf blade anatomy.•Cross-sections usually with 7 vascular bundles, 5 ribs above; sclerenchyma under abaxial epidermis discontinuous, forming large fascicles, adaxial sclerenchyma present and extending to every second vascular bundle; adaxial epidermis with scattered hairs, the hairs ca. 0.1 mm long. Observations. This Venezuelan species is geographically isolated and known only from the type collection. Festuca turimiquirensis differs from F. tolucensis by having smaller spikelets (lower glumes 3-4 versus 5-8.5 mm long, upper glumes 4•5.5 versus 5-9.5 mm long), opened panicles (6-7 versus 0.5-1.5 cm wide), and the abaxial sclerenchyma is discontinuous (versus continuous). Distribution and habitat.•Festuca turimiquirensis is restricted to the Serrania de Turimiquire (Anzoategui) of Venezuela where it occurs in grasslands on rocky slopes in open areas between 2000-2350 m. 56. Festuca vaginalis (Benth.) Laegaard, Novon 8(1): 30. 1988. (Figs. 64, 71, 97D). Poa vaginalis Benth., PI. Hartw. 261. 1846. TYPE: Ecuador. Pichincha, Hacienda de Antisana, K.T. Hartweg 1450 (holotype: K!; isotype: US-88715!). 143 Tussocked perennials with intravaginal innovations. Culms 20-60 cm tall, erect, glabrous; nodes 1, basal and 1 or 2 short leaves. Leaf sheaths coriaceous, stramineous, occasionally grayish, glabrous; ligules 0.8-1.2 mm long, coriaceous, oblong or triangular, apex short-ciliate; blades (4•)7•17 cm long, 0.4-0.8 mm wide, conduplicate, glaucous. Panicles 10-15 x 0.5 cm, contracted, narrow, whitish or purplish-white, occasionally pale green; Spikelets 8-9.5 mm long, lanceolate, florets 2 or 3; rachilla short-pubescent; glumes (5.5-)6.5-9 mm long, almost as long as the spikelet, membranous, white or with purple stripes, midnerve scabrous distally, apex obtuse; lower glumes (5.5•)6.5•8 (-9) mm long, 1-nerved; upper glumes 6.5-9 mm long, 3-nerved; lemmas 7.5-8.5 mm long, oblong, 5-nerved, membranous, apex short two-dentate, awned, the awn 0.5-1 mm long; callus glabrous; paleas 3/4 as long as the lemma, scabrous on keels; lodicules lanceolate; anthers 0.6-0.9 mm long; ovary apex glabrous. Caryopses lanceolate, hilum 2/3 as long as the grain, linear. Leaf blade anatomy.•Cross-sections with (7•)9•11 vascular bundles and 5-9 ribs above; sclerenchyma discontinuous under both abaxial and adaxial epidermis, girders absent; bulliform cells absent; adaxial epidermis with scattered hairs, the hairs 0.075-0.09 mm long. Observations.•Festuca vaginalis is morphologically distinct from the other species treated in this revision and we do not know which species might be its closest sister. Festuca vaginalis has been mentioned as occurring in Ecuador and Colombia (Luteyn 1999, Rangel 2000); although its occurrence in Colombia has not been confirmed. Distribution and habitat.• Festuca vaginalis is endemic to Ecuador. It occurs on volcanic rocks in superparamos and dry grass paramos between 3500-5000 m. It also has spread secondarily onto disturbed areas of dry grass paramos and along road embankments, so called "arenal." KEY TO THE SUBSPECIES OF FESTUCA VAGINALIS la. Culms 30-60 cm long, robust; leaf blades ca. 0.8 mm wide; panicles whitish or purplish-white.... 56a. F. vaginalis subsp. vaginalis lb. Culms 20-30 cm tall, not robust; leaf blades 0.4-0.5(-0.6) mm wide; panicles pale-green 56b. F. vaginalis subsp. cayambae 144 Festuca in South American Paramos Figure 71. Festuca vaginalis subsp. vaginalis A. & B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 3134 (COL). Festuca in South American Paramos 56A. Festuca vaginalis subsp. vaginalis (Fig. 71). Culms 30-60 cm long, robust. Leaf blades ca. 0.8 mm wide. Panicles whitish or purplish-white. Leaf blade anatomy.•Cross-sections with 9-11 vascular bundles. Additional specimens examined. ECUADOR. Bolivar: Volcan Chimborazo, W side, 1°28'S, 78°48'W, 4800 m, 14 Sep 1995, Sklendf & Kosteckovd 143-2 (QCNE); road Salinas-Los Arenales, km 6, 01°22'S, 79°00'W, 4000 m, S. Laegaard 55327 (AAU); valley 4 km S of Cruces de los Arenales, 4270^1300 m, 3 Mar 1992, S. Laegaard 101559 (AAU, QCA); 01°25'S, 78°54'W, 4160-4200 m, S. Laegaard 101495 (AAU); km 66.5 SW of Ambato on Hwy to Guaranda and 1.2 on road Fecundo Vela, km 1.2 km, 4290 m, 03 May 1990, P.M. Peterson 8987 & C.R. Annable (K, MO, US, QCNE). Cotopaxi: Angamarca road, km 5 from junction to Latacunga-La Mana road, 01°00'S, 78°55'W, 4250-4280 m, S. Laegaard 102149 (AAU); Volcan Cotopaxi, road to Refugium, 00°40'S, 78°26'W, 4350 m, S. Laegaard 103127 (AAU); SW slope, 4100 m, E. Asplund 7497 (F, K, NY, S, US); El Tambo SE of Volcan Cotopaxi, 00°42'S, 78°18'W, 3650 m, S. Laegaard 55536 (AAU); Hacienda Pauzacha, 00°44'S, 78°29'W, 3650 m, S. Laegaard 55735 (AAU); road Zumbahua-Pujili, km 39, 00°53'S, 78°47'W, 3750-3800 m, S. Laegaard 102096 (AAU); Parque Nacional Cotopaxi, Laguna Limpiopungo, 3800 m, P.M. Peterson 8739, C.R. Annable & M.E. Poston (MO, QCA, QCNE, US); plate below NE side of volcano, 00°37'S, 78°24'W, 3800 m, 26 Jun 1999, D. Stancik 3137 (PRC, QCA); NE side of volcano Cotopaxi, 00°37.5'S, 78°24.2'W, 3820 m, 26 Jun 1999, D. Stancik 3134 (PRC, QCA); valley NW of Limpiopungo to foot of Volcan Ruminahui, 00°38'S, 78°28'W, 4300 m, 25 Feb 1992, S. Laegaard 101460 (AAU, QCNE). Laguna Quilotoa, 00°51'S, 78°53'W, 3480-3820 m, S. Laegaard 101335A (AAU); Panamerican Hwy, km 2 S of Cruce de Cordillera, 00°39'S, 78°35'W, 3500 m, 13 Aug 1984, S. Laegaard 52656 (AAU, QCNE); S. Laegaard 52659 (AAU); S of Cruce de Arenales, km 5, 01°32'S, 78°52'W, 3950 m, 18 Sep 1998, S Laegaard 19159 (AAU, LOJA); Rancho Santa Maria, 00°43'S, 78°29'W, 3750-3950 m, S. Laegaard 52202 (AAU); Mun. Chaupi, NE slope of volcan Illiniza Norte, 00°38'11"S, 78°43T'W, 4450 m, 12 Oct 2000, D. Stancik 4013 (PRC, QCA); SE slope of volcan, 00°38'45"S, 78°25'32"W, 145 4300 m, 28 Sep 2000, D. Stancik 3881 (PRC, QCA, US). Chimborazo: Volcan Chimborazo, lower Refugio, 01°28'S, 78°50'W, 4840 m, S Laegaard 102796 (AAU,QCNE); 5000 m, 11 May 1992, Rauh & Hirsch E315 (US); road from Refugium, km 4.7, 01°27'S, 78°52'W, 4330^1400 m, S Laegaard 102795 (AAU, QCNE); Chimborazo, Sodiro s.n. (US); lower Refugium, 01°27'S, 78°50'W, 4800 m, 5 Mar 1988, S. Laegaard et al. 70553 (QCA); Sodiro s.n. (QPLS); 01°27'S, 78°50'W, 4800 m, S. Laegaard et al. 70554 (AAU, QCNE); paroquilla San Juan, 01°30'S, 78°49'W, 4150 m, 4 Aug 1992, Ceron et al. 19822 (QAP); Cruce de los Arenales, km 5, 01°27'S, 78°52'W, 4330^1400 m, S. Laegaard 102795 (AAU, QCNE); km 4.7 from lower refugio of Volcano Chimborazo, arenales, 01°27'S, 78°50'W, 4500 m, 5 Mar 1988, S Laegaard & S.A. Renvoize 70558 (K); Panamerican Hwy. RiobambaAmbato,km22,01°30'S,78°42'W, 3500 m, 11 Oct 1985, S Laegaard 55400 (AAU, QCA); S side of Volcano, 4300 m, 1929, Rimbach 97a (US); Urbina - paramo on E flank of Mt. Chimborazo, 3600 m, A.S Hitchcock22019 (K, NY, US); S slope, 3800 m, Fagerlind& Wibom 934bis (S); Fagerlind & Wibom 932 (S); Dec 1890, Sodiro s.n. (QPLS); 4600 m, E. Asplund 8399 (S); Nudo delgualata-Sanancajas, 3600 m, Acosta-Solis 21187 (US); km 18.2 ESE of Lago Colta on road to Pallatanga, 3700 m, 21 May 1990, P.M. Peterson 9228, E.J. Judziewicz, R.M. King & P.M. Jorgensen (MO, QCA, QCNE, US); Prov. Riobamba, praed Toledo, Sep 1891, Sodiro s.n. (P); Mun. Guaranda, W side of volcan Chimborazo, road to Refugium, 0 P28 'S, 78°52'W, 4100 m, 5 Jul 1999, D. Stancik 3163 (PRC, QCA); Mun. Riobamba, volcan Chimborazo, around upper Refugio, 5000 m, 20 Sep 2000, D. Stancik 3706 (PRC, QCA); sector Cruce de los Arenales, 01°28'15"S, 78°54'6" W, 4300 m, 20 Sep 2000, D. Stancik3718 (PRC, QCA). Imbabura: Paramo on the N side of Nevado Cayambe, 0°03'07"N, 77°59'27"W, 17 Aug 2004, Sklendf 8067, 8115 (PRC). MoronaSantiago: Hda. Hargualla-Hda. San Eduardo, way to Parque Nacional Sangay, 02°0.25'S, 78°27'W, 3700 m, 19 Jul 1999, D. Stancik 3323 (PRC, QCA); Parque Nacional Sangay, a confluence of Quebrada Plazabamba Chico with Q. Plazabamba, 02°0.48'S, 78°25.4'W, 3600 m, 21 Jul 1999, D. Stancik 3341 (PRC, QCA). Pichincha: Cerro Antisana, W. Huagrahialina Campamento, 4200 m, Black 278, 279 (AAU); 4600 m, E. Asplund 17338 (NY, S); 00°32'S, 78°12'W, type locality, 4030 m, S. Laegaard 102848 (AAU); between Antisana and 146 Festuca in South American Paramos Sincholagua, E. Asplund 3648 (US); Rucu Pichincha, NE slope, 00°10'S, 78°34'W, 4300^1500 m, Sklendf & Kosteckovd 198 (AAU); Lloa-Guagua Pichinha road, km 11, 00°12'S, 78°35'W, 4310 m, 9 May 1992, S. Laegaard 102743 (AAU, QCNE); around Refugium, 00°12'S, 78°35'W, 4550 m, S. Laegaard 102 758 (AAU); 4600 m, E. Asplund 7397 (K, NY, S, US); Pichincha, Cotocollao, Sodiro s.n. (QPLS); 3200 m, E. Asplund 6620 (S, US); Paramo de Guamani, 00°18'S, 78°14'W, 3750 m, S. Laegaard 51386 (AAU, QCNE); 00°20'S, 78°14'W, 3700 m, S. Laegaard 19598 (AAU, LOJA, PRC, QCNE); 3600-3800 m, E. Asplund 8721 (S); 4050 m, Barclay & Juajibioy 8905 (COL, US); Andium Quitensium, Jameson 230 (P); Mun. Pifo, Paramo de Guamani, 00°20'S, 78°12'W, 4000 m, 19 Jun 1999, D. Stancik 3014 (PRC, QCA); 3700 m, D. Stancik 3020 (PRC, QCA, US); D. Stancik 3001 (PRC, QCA). Tungurahua: Volcan Chimborazo, SW of San Fernando, 3500 m, 23 Jul 1959, Barclay & Juajibioy 8166 (COL, US); above Mocha, 3300 m, E. Asplund 7872 (S). Additional specimens examined. ECUADOR. Bolivar: km 60.4 SW of Ambato on Hwy to Guaranda, 4090 m, P.M. Peterson & C.R. Annable 8984 (QCNE, US); 01°25'S, 78°52'W, 3950 m, 15 Feb 1999, S. Laegaard 19589 (AAU, PRC); km 66.5, 3760 m, 3 May 1990, P.M. Peterson & C.R. Annable <WW(K,MO,QCA,US);km33,01°18'S,78°48'W, 3700 m, 1 Mar 1992, S Laegaard 101505 (AAU, QCNE); Cruce de los Arenales, 01°25'S, 78°54'W, 4160-4200 m, S. Laegaard 101493 (AAU, PRC). Caflar: Panamerican Hwy. ca. 5 km S of Canar, 02°33'S,78°56'W, 3500 m, S Laegaard 105123 (AAU); 3350 m, S Laegaard 101828 (AAU). Chimborazo: Mun. Guaranda, pasture around Vereda Rio Colorado, 4200 m, 2 Jul 1999, D. Stancik 3182 (PRC, QCA, W). Cotopaxi: Mun. Chaupi, NE slope of volcan Illiniza Norte, 00°37'59"S, 78°42'42"W, 4000-4050 m, 12 Oct 2000, D. Stancik 4033, 4035 (PRC, QCA). Pichincha: Hacienda Antisana and Mt. Sincholagua, E. Asplund 8648 (S). 56b. Festuca vaginalis subsp. cayambae Stancik, Folia Geobot. Phytotax. 39(1): 107. 2004. TYPE: Ecuador. Pichincha, along road to Volcan Cayambe, near entrance to National Park, 78°04'W, 00°03'S, 3550 m, 1 Mar 1988, S. Laegaard & S.A. Renvoize 70488 (holotype: AAU!; isotypes: QCA!, QCNE!, K!). Festuca sublimis forma vivipara St.-Yves, Candollea 3: 255. 1927. Type: Peru. Puca de Ladrillos, entre Pacasmayo et Mayobamba, 3400 m, Stiibel Fl. Peru 30 (syntype: B?); Ecuador. Ilizna, Tisiche, Stiibel Fl. Ecuador 301 (syntype: B?). Culms 20-30 cm tall, not robust. Leaf blades 0.4-0.5(-0.6) mm wide. Panicles pale-green. Leaf blade anatomy.•Cross-sections with 7 vascular bundles. EXCLUDED NAME Comments.•The status of this taxon is in question since we were unable to locate any of the syntypes. Alexeev (1986) cited this taxon as a synonym of F asplundii but did not indicate that he saw any of the original material. SCANNING ELECTRON MICROGRAPHS OF LEAF EPIDERMAL SURFACES Figure 72. Leaf blade surfaces. A-D. Festuca amplissima subsp. amplissima. A. Abaxial epidermis with ribs, densely covered with prickles. B. Abaxial epidermis, detail view of long cell with sinuous walls, rounded silica bodies, and prickles. C. Adaxial epidermis with long cells and regularly occurring stomata. D. Adaxial epidermis with long and short (silica bodies) cells, stomata (surrounded by wax), and prickles on the margin. E & F. F. coromotensis. E. Abaxial epidermis with regular ribs. F. Abaxial epidermis with detail of oval (sinuous) silica bodies and prickles. A & B, Barclay 7033 (COL); C & D, Liebmann 6110 (C); E, F, Stancik 4180 (PRC). 148 Festuca in South American Paramos Figure 73. Leaf blade surfaces. A & B. Festuca coromotensis. A. Adaxial epidermis with ribs, sparsely covered with short macrohairs. B. Adaxial epidermis, detail view of silica bodies, stomata and macro-hairs. C-F. F. elviae. C. Abaxial epidermis with long cells and regularly occurring silica bodies. D. Abaxial epidermis, detail view of silica bodies with short cork cell. E. Adaxial epidermis with regular ribs and short macro-hairs. F. Adaxial epidermis, detail view of stomata and cell walls covered by fine crystals of wax. A & B, Stancik 4180 (PRC); C-E, Stancik 4178 (PRC); F, Stancik 4171 (PRC). Festuca in South American Paramos 149 Figure 74. Leaf blade surfaces. A-D. Festuca flacca. A. Abaxial epidermis with small ribs densely covered with prickles. B. Adaxial epidermis covered with prickles and stomata. C. Adaxial epidermis, detail view of stoma and prickle. D. Adaxial epidermis, detail view of wax crystals. E & F. F. guaramacalana. E. Abaxial epidermis with regular ribs F. Abaxial epidermis with detail of circular silica bodies. A-D, Stancik 4180 (PRC); E & F, Stancik 4286 (PRC). 150 Festuca in South American Paramos Figure 75. Leaf blade surfaces. A & B. Festuca guaramacalana. A. Adaxial epidermis with ribs, covered with macro-hairs. B. Adaxial epidermis, detail view of macro-hairs. C-F. F. sodiroana. C. Abaxial epidermis with small ribs and occasionally occurring prickles. D. Abaxial epidermis, detail view of long and short (silica bodies) cells. E. Adaxial epidermis with regular ribs and sparse short macro-hairs. F. Adaxial epidermis, detail view of stoma. A & B, Stancik 4286 (PRC); C-F, Stancik 2632 (PRC). Festuca in South American Paramos 151 I'll I I F •ft 50 urn "••'"''-""•'•'• • Figure 76. Leaf blade surfaces. A-D. Festuca ulochaeta. A. Abaxial epidermis with small ribs. B. Abaxial epidermis, detail view of prickle and silica bodies with cork cells. C. Adaxial epidermis with small ribs and regurally occurring stomata and short macrohairs. D. Adaxial epidermis, detail view of stomata and macro-hairs. E & F. F. caldasii. E. Abaxial epidermis densely covered with prickles. F. Abaxial epidermis with detail of oval (sinuous) silica bodies and prickles. A-D, Stancik 4179 (PRC); E & F, Laegaard 20405 (AAU). 152 Festuca in South American Paramos Figure 77. Leaf blade surfaces. A & B. Festuca caldasii. A. Adaxial epidermis with small ribs and regurally distributed stomata and macro-hairs. B. Adaxial epidermis, detail view of stomata covered by wax. C-F. F. reclinata. C. Abaxial epidermis with long cells and regularly occurring stomata and slender prickles. D. Abaxial epidermis, detail view of prickles and stomata. E. Adaxial epidermis with small ribs; regurally occurring prickles and stomata. F. Adaxial epidermis with detail of prickles and wax covering cell walls. A & B, Laegaard 102535 (AAU); C-F, Cuatrecasas & Barriga 9970 (COL). Festuca in South American Paramos 153 Figure 78. Leaf blade surfaces. A-D. Festuca woodii. A. Abaxial epidermis with ribs, densely covered with prickles. B. Adaxial epidermis with pronounced ribs densely covered with prickles. C. Adaxial epidermis with detail view of stomata laying between two ribs. D. Adaxial epidermis with detail view of prickles and waxes covering cell walls. E & F. F. arundinacea. E. Abaxial epidermis with regular ribs and rows of prickles. F. Abaxial epidermis with detail view of stomata and silica bodies. A-D, Wood 5254 (COL); E & F, Stancik3221 (PRC). 154 Festuca in South American Paramos Figure 79. Leaf blade surfaces. A & B. Festuca arundinacea. A. Adaxial epidermis with rows of stomata between small ribs. B. Adaxial epidermis, detail view of long cells interrupted by short cells (silica bodies and cork cells). C-F. F. quadridentata. C. Abaxial epidermis covered by prickles. D. Abaxial epidermis with detail view of prickles and short cells. E. Adaxial epidermis with pronounced ribs densely covered with prickles and wax. F. Adaxial epidermis with detail of wax and prickles. A & B, Stancik 3221 (PRC); C-F, Laegaard & Sklendf 20308 (PRC). Festuca in South American Paramos 155 Figure 80. Leaf blade surfaces. A. Festuca quadridentata. Abaxial epidermis with detail view of wax. B-F. F. venezuelana. B. Abaxial epidermis with small ribs covered by prickles. C. Abaxial epidermis with detail view of silica bodies and prickles. D. Adaxial epidermis with small ribs with prickles, stomata and silica bodies. E. Adaxial epidermis. F. Adaxial epidermis with detail view of prickles, stomata and silica bodies. A, Laegaard & Sklendf 20308 (PRC); B-F, Stancik 4263(PRC). 156 Festuca in South American Paramos 50 (Jin Figure 81. Leaf blade surfaces. A-F. Festuca fragilis. A. Abaxial epidermis densely covered with prickles and silica bodies. B, C. Abaxial epidermis with detail view of silica bodies and prickles. D. Adaxial epidermis with ribs and macro-hairs. Stomata on the bottom of ribs. E. Adaxial epidermis with detail view of macro-hairs and stomata. F. Adaxial epidermis with detail view of wax covering cell walls. A-C, Wood 5259 (COL); D-F, Stancik4248 (PRC). Festuca in South American Paramos 157 Figure 82. Leaf blade surfaces. A-E. Festuca fimbriata. A. Abaxial epidermis with ribs. B. Abaxial epidermis, detail view of silica bodies, prickles and stomata. C. Adaxial epidermis with pronounced ribs covered densely with prickles. D. Adaxial epidermis, detail view of ribs with prickles and silica bodies on the top of ribs, stomata between the ribs. E. Adaxial epidermis, detail view of stoma and wax on the cell walls. F. F. rubra. Abaxial epidermis with long and short cells. A-E, Brade & Kuhlman 15631 (PRC); F, Stancik 3457 (PRC). Festuca in South American Paramos 158 E 8^1 : w ,.\L. . ^^H J, 1 IP Hw wA aih 9^ L100VU I Figure 83. Leaf blade surfaces. A & B. Festuca rubra. A. Adaxial epidermis with ribs covered with macro-hairs. B. Adaxial epidermis, detail view of macro-hairs and stomata on the base of the ribs. C & D. F. andicola. C. Abaxial epidermis of the involute blade with macro-hairs on the margin. D. Adaxial epidermis, detail view of macro-hair and wax on the cell walls. E & F. F. soukupii. E. Abaxial epidermis of the involute blade with silica bodies and macro-hairs on the margin. F. Adaxial epidermis with macro-hairs on the ribs. A & B, Stancik 3457 (PRC); C & D, Laegaard 52807 (AAU); E &F, Stancik 3455 (PRC). Festuca in South American Paramos 159 Figure 84. Leaf blade surfaces. A-D. Festuca azucarica. A. Abaxial epidermis with long and short cells. B. Abaxial epidermis, detail view of rounded silica bodies. C. Adaxial epidermis with ribs covered with long macro-hairs. D. Adaxial epidermis, detail view of stomata at the bottom of rib (middle part of this image). E & F. F. colombiana. E. Abaxial epidermis with long and short cells. F. Adaxial epidermis, macro-hairs on the lateral walls of the ribs. A-D, Stancik 3405 (PRC); E & F, Stancik 2561 (PRC). 160 Festuca in South American Paramos 1 WHm "f Figure 85. Leaf blade surfaces. A & B. Festuca colombiana. A. Adaxial epidermis, detail view of stomata and macro-hairs. B. Adaxial epidermis, detail view of wax. C & D F. dasyantha. C. Abaxial epidermis with small ribs, long and short cells. D. Adaxial epidermis with ribs covered with long macro-hairs. E & F. F. hatico. E. Abaxial epidermis with small ribs. F. Abaxial epidermis, detail view of silica bodies. A & B, Stancik 1289 (PRC); C & D, Laegaard 101260 (AAU); E & F, Stancik 4289 (PRC). Festuca in South American Paramos 161 Figure 86. Leaf blade surfaces. A & B. Festuca hatico. A. Adaxial epidermis with ribs, densely covered with macro-hairs. B. Adaxial epidermis, detail view of macro-hairs and wax deposits. C-F. F. laegaardii. C. Abaxial epidermis with small ribs. D. Abaxial epidermis, detail view of long and short cells (primarily silica bodies). E. Adaxial epidermis with ribs covered with macro-hairs, stomata on the base of the ribs. F. Adaxial epidermis, detail view of stomata. A & B, Stancik 4289 (PRC); C-F, Stancik 3365 (PRC). 162 Festuca in South American Paramos Figure 87. Leaf blade surfaces. A & B. Festuca pilar-franceii. A. Abaxial epidermis, detail view of silica bodies. B. Adaxial epidermis, detail view of ribs covered with long macro-hairs. C-F. F. procera. C. Abaxial epidermis with small ribs. D. Abaxial epidermis with long and short (silica bodies) cells. E. Adaxial epidermis with ribs covered by long macro-hairs. F. Adaxial epidermis, detail view of macro-hair and wax deposits. A & B, Stancik 241 (PRC); C-F, Stancik 4106 (PRC). Festuca in South American Paramos 163 . c \ RJI ti^l 1 ~-y- 50 jjm Figure 88. Leaf blade surfaces. A & B. Festuca toca. A. Abaxial epidermis with small rib and silica bodies. B. Adaxial epidermis, detail view of rib covered with macro-hairs, stomata are on the base of the rib. C-F. F. asplundii. C. Abaxial epidermis sparsely covered with prickles. D. Abaxial epidermis, detail view of prickles and silica bodies. E. Adaxial epidermis with ribs covered with macro-hairs. F. Adaxial epidermis, detail view of macro-hairs and stomata on the base of the rib. A & B, Stancik 1418 (PRC); C-F, Stancik 3611 (PRC). 164 Festuca in South American Paramos >_-^J- -FV-^ * ; •::-. .; r - r ' ** • i . •- 1 ^ ,.. -• J • I'' 1 >% 1 1 r " ' * 50 (jm Figure 89. Leaf blade surfaces. A & B. Festuca boyacensis. A. Abaxial epidermis, detail view of silica bodies. B. Adaxial epidermis, detail view of stomata and macro-hairs and cell walls covered with wax. C & D. F. carchiense. C. Abaxial epidermis, view of long and short cells and sparse prickles. D. Adaxial epidermis with ribs covered with short macro-hairs. E & F. F. chimborazensis subsp. chimborazensis. E. Abaxial epidermis with small ribs. F. Abaxial epidermis, detail view of silica bodies. A & B, Stancik 2166 (PRC); C & D, Laegaard 101716 (AAU); E & F, Stancik 4033B (PRC). Festuca in South American Paramos 165 L ! * 1 - 20Mm]| Figure 90. Leaf blade surfaces. A & B. Festuca chimborazensis. A. Adaxial epidermis with ribs with stomata between long cells, sparsely covered with short macro-hairs. B. Adaxial epidermis, detail view of prickles (margin of the leaves), macro-hairs and stomata. C-F. Festuca cleefiana. C. Abaxial epidermis. D. Abaxial epidermis, detail view of silica bodies. E. Adaxial epidermis with ribs covered with macro-hairs. F. Adaxial epidermis, detail view of stomata on the base of the ribs. A & B, Stancik 4033B (PRC); C-F, Stancik 2020 (PRC). 166 Festuca in South American Paramos Figure 91. Leaf blade surfaces. A-D. Festuca cocuyana. A. Abaxial epidermis with long and short cells. B. Abaxial epidermis, detail view of silica bodies. C. Adaxial epidermis with several ribs covered with macro-hairs, sometimes with silica-bodies or stomata between long cells. D. Adaxial epidermis with detail view of macro-hairs, stomata and wax. E & F. F. cundinamarcae. E. Abaxial epidermis. F. Abaxial epidermis with detail view of silica bodies. A-D, Cleef9078 (COL); E & F, Stancik 2114 (PRC). Festuca in South American Paramos 167 Figure 92. Leaf blade surfaces. A & B. Festuca cundinamarceae. A. Adaxial epidermis with ribs densely covered with macro-hairs. B. Adaxial epidermis, detail view of stomata at the base of the ribs. C-F. F. densipaniculata. C. Abaxial epidermis densely covered with prickles. D. Abaxial epidermis with detail view of prickles and wax. E. Adaxial epidermis with regular ribs covered with macro-hairs. Stomata on the base of ribs (right margin of the image). F. Adaxial epidermis with detail of macro-hairs and stomata. A & B, Stancik 2114 (PRC); C-F, Asplund 8397 (S). 168 Festuca in South American Paramos B • • , 100 pm Figure 93. Leaf blade surfaces. A-D. Festuca glumosa. A. Abaxial epidermis of the involute leaf with rows of prickles. B. Abaxial epidermis, detail view of silica bodies and prickles. C. Adaxial epidermis with ribs covered with short macro-hairs. D. Adaxial epidermis with detail view of macro-hairs and stomata. Cell walls covered with wax. E & F. F. glyceriantha. E. Adaxial epidermis with regular ribs covered with macro-hairs. F. Adaxial epidermis with detail of wax. A-D, Stancik 4180 (PRC); E & F, Peterson 14936 (US). Festuca in South American Paramos 169 Figure 94. Leaf blade surfaces. A & B. Festuca imbaburensis. A. Adaxial epidermis with ribs covered with macro-hairs. Cell walls covered with wax deposits. B. Adaxial epidermis, detail view of wax, stomata and macro-hair. C-F. F. monguensis. C. Abaxial epidermis of the involute leaf, with small ribs and prickles on the margin. D. Adaxial epidermis with detail view of prickles and silica bodies. E. Adaxial epidermis with regular ribs covered with macro-hairs. F. Adaxial epidermis with detail of macro-hairs and stomata (at the bottom of ribs). A & B, Stancik 4098 (AAU); C-F, Stancik 2025 (PRC). 170 Festuca in South American Paramos Figure 95. Leaf blade surfaces. A-D. Festuca nereidaensis. A. Abaxial epidermis with long and short cells. B. Abaxial epidermis, detail view of silica bodies. C. Adaxial epidermis with ribs covered densely with macro-hairs. D. Adaxial epidermis with detail view of macro-hairs. Stomata on the bottom of the ribs (not seen). E & F. F. oroana. E. Adaxial epidermis with regular ribs covered densely with macro-hairs. F. Adaxial epidermis with detail view of macro-hairs and dense coverage of wax. A-D, Stancik 340IB (PRC); E & F, Laegaard 70309 (AAU). Festuca in South American Paramos 171 Figure 96. Leaf blade surfaces. A-D. Festuca sanctae-martae. A. Abaxial epidermis with small ribs and prominent long and short cells. B. Abaxial epidermis, detail view of silica bodies. C. Adaxial epidermis with small ribs and short macro-hairs. D. Adaxial epidermis with detail view of macro-hairs and stomata between long cells. E & F. F. subulifolia. E. Abaxial epidermis with detail view of prickles and silica bodies. F. Adaxial epidermis with small ribs covered with macro-hairs and stomata at the bottom between ribs. A-D, Cuatrecasas & Castaneda 24532 (COL); E & F, Stancik 2914 (PRC). 172 Festuca in South American Paramos Figure 97. Leaf blade surfaces. A-C. Festuca tolucensis subsp. tolucensis. A. Abaxial epidermis densely covered with prickles and silica bodies. B. Adaxial epidermis with ribs covered with macro-hairs, stomata at the bottom between the ribs. C. Adaxial epidermis, detail view of stomata and macro-hairs. D. F. vaginalis. Adaxial epidermis with ribs covered with short macro-hairs and deposits of wax on the cell walls. A-C, Stancik 4279 (PRC); D, Laegaard 101559 (AAU). Festuca in South American Paramos 173 Appendices INDEX TO FIGURES Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. 1. Illustration of Festuca amplissima 2. Distribution map of Festuca amplissima, F coromotensis, and Fflacca 3. Illustration of Festuca coromotensis 4. Illustration of Festuca elviae 5. Illustration of Festuca flacca 6. Distribution map of Festuca sodiroana, F elviae, and F guaramacalana 7. Illustration of Festuca guaramacalana 8. Illustration of Festuca sodiroana 9. Illustration of Festuca tovariensis 10. Illustration of Festuca ulochaeta 11. Distribution map of Festuca ulochaeta, F tovariensis, and F caldasii 12. Illustration of Festuca caldasii 13. Illustration of Festuca reclinata 14. Distribution map of Festuca pratensis, F woodii, andF reclinata 15. Illustration of Festuca woo dii 16. Illustration of Festuca arundinaceae 17. Distribution map of Festuca dichoclada, F arundinacea, and F. venezuelana 18. Illustration of Festuca quadridentata 19. Illustration of Festuca venezuelana 20. Illustration of Festuca fragilis 21. Distribution map of Festuca quadridentata, F fragilis, andF rubra 22. Illustration of Festuca fimbriata 23. Distribution map ofFestuca fimbriata, andF andicola 24. Illustration of Festuca andicola 25. Illustration of Festuca rubra 26. Illustration of Festuca soukupii 27. Distribution map of Festuca soukupii, F chitagana, and F hatico 28. Illustration of Festuca azucarica 29. Distribution map of Festuca colombiana, F dasyantha, F azucarica, and F laegaardi 30. Illustration of Festuca chitagana 31. Illustration of Festuca colombiana 32. Illustration of Festuca dasyantha 33. Illustration of Festuca hatico 34. Illustration of Festuca laegaardii 35. Illustration of Festuca pilar-franceii 36. Illustration of Festuca procera 3 7. Distribution map of Festuca procera, F pilar-franceii, and F boyacensis 38. Illustration of Festuca toca 39. Distribution map of Festuca asplundii, F toca, and F cocuyana 40. Illustration of Festuca asplundii 41. Illustration of Festuca boyacensis 42. Illustration of Festuca carchiense 43. Illustration of Festuca chimborazensis 44. Distribution map of Festuca chimborazensis, F carchiense, F cleefiana, and F dinirica 45. Illustration of Festuca chita 46. Illustration of Festuca cleefiana 47. Illustration of Festuca cocuyana 19 20 21 23 25 26 27 29 32 34 35 37 39 40 41 43 45 47 49 50 51 53 54 56 58 60 61 64 65 67 68 71 73 74 76 78 79 82 83 85 89 90 92 93 96 97 99 174 Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Festuca in South American Paramos 48. 49. 50. 51. 52. Illustration of Festuca cundinamarcae 101 Distribution map of Festuca glumosa, F cundinamarcae, F chita, and F. turimiquirensis.... 102 Illustration of Festuca densipaniculata 103 Illustration of Festuca dinirica 105 Distribution map of Festuca glyceriantha, F holubii, F densipaniculata, and F monguensis .... 106 53. Illustration of Festuca glumosa 107 54. Illustration of Festuca glyceriantha 110 55. Illustration of Festuca holubii 112 56. Illustration of Festuca huamachensis 114 57. Distribution map of Festuca huamachucensis, F imbaburensis, F nereidaensis, and F sanctaemartae 115 58. Illustration of Festuca imbaburensis 116 59. Illustration of Festuca monguensis 118 60. Illustration of Festuca nereidaensis 120 61. Illustration of Festuca oroana 121 62. Illustration of Festuca parciflora 123 63. Illustration of Festuca renvoizei 125 64. Distribution map of Festuca vaginalis, F parciflora, F oroana, and F renvoizii 126 65. Illustration of Festuca sanctae-martae 127 66. Illustration of Festuca subulifolia 128 67. Distribution map of Festuca tolucensis, F subulifolia,and F sumapana 129 68. Illustration of Festuca sumapana 137 69. Illustration of Festuca tolucensis 138 70. Illustration of Festuca turimiquirensis 142 71. Illustration of Festuca vaginalis 144 72. Leaf blade surfaces of Festuca amplissima and F coromotensis 147 73. Leaf blade surfaces of Festuca coromotensis and F elviae 148 74. Leaf blade surfaces of Festuca flacca and F guaramacalana 149 75. Leaf blade surfaces of Festuca guaramacalana and F sodiroana 150 76. Leaf blade surfaces of Festuca ulochaeta andF caldasii 151 77. Leaf blade surfaces of Festuca caldasii and F reclinata 152 78. Leaf blade surfaces of Festuca woodii andF arundinacea 153 79. Leaf blade surfaces of Festuca arundinacea and F quadridentata 154 80. Leaf blade surfaces of Festuca quadridentata andF venezuelana 155 81. Leaf blade surfaces of Festuca fragilis 156 82. Leaf blade surfaces of Festuca fimbriata andF rubra 157 83. Leaf blade surfaces of Festuca rubra, F andicola, andF soukupii 158 84. Leaf blade surfaces of Festuca azucarica andF colombiana 159 85. Leaf blade surfaces of Festuca colombiana, F. dasyantha, andF hatico 160 86. Leaf blade surfaces of Festuca hatico andF laegaardii 161 87. Leaf blade surfaces of Festuca pilar-franceii and F procera 162 88. Leaf blade surfaces of Festuca toca andF asplundii 163 89. Leaf blade surfaces of Festuca boyacensis, F carchiense, andF chimborazensis 164 90. Leaf blade surfaces of Festuca chimborazensis and Festuca cleefiana 165 91. Leaf blade surfaces of Festuca cocuyana andF cundinamarcae 166 92. Leaf blade surfaces of Festuca cundinamarceae andF densipaniculata 167 93. Leaf blade surfaces of Festuca glumosa and F glyceriantha 168 94. Leaf blade surfaces of Festuca imbaburensis andF monguensis 169 95. Leaf blade surfaces of Festuca nereidaensis andF oroana 170 96. Leaf blade surfaces of Festuca sanctae-martae andF subulifolia 171 97. Leaf blade surfaces of Festuca tolucensis andF vaginalis 172 Festuca in South American Paramos ALPHABETICAL LIST OF ACCEPTED SPECIES AND SUBSPECIES Number in parentheses corresponds to the number of the species in the taxonomic treatment. uca amplissima subsp. amplissima (la) uca amplissima subsp. sierrae (lb) uca andicola (19) uca arundinacea (12) uca asplundii (31) uca azucarica (22) uca boyacensis (32) uca caldasii (9) uca carchiense (33) uca chimborazensis subsp. chimborazensis (34a) uca chimborazensis subsp. micacochensis (34b) uca chita (35) uca chitagana (23) uca cleefiana (36) uca cocuyana (37) uca colombiana (24) uca coromotensis (2) uca cundinamarcae (38) uca dasyantha (25) uca densipaniculata (39) uca dichoclada (14) uca dinirica (40) uca elviae (3) ucafimbriata (18) ucaflacca (4) ucafragilis (17) ucaglumosa (41) wca glyceriantha (42) wca guaramacalana (5) wca hatico (26) Festuca holubii (43) Festuca huamachucensis (44) Festuca imbaburensis (45) Festuca laegaardii (27) Festuca monguensis (46) Festuca nereidaensis (47) Festuca oroana (48) Festuca parciflora subsp. loxana (49b) Festuca parciflora subsp. parciflora (49a) Festuca pilar-franceii (28) Festuca procera (29) Festuca pratensis (13) Festuca quadridentata (15) Festuca reclinata (10) Festuca renvoizei (50) Festuca rubra (20) Festuca sanctae-martae (51) Festuca sodiroana (6) Festuca soukupii (21) Festuca subulifolia (52) Festuca sumapana (53) Festuca toca (30) Festuca tolucensis subsp. culata (54c) Festuca tolucensis subsp. perijae (54b) Festuca tolucensis subsp. tolucensis (54a) Festuca tovariensis (7) Festuca turimiquirensis (55) Festuca ulochaeta (8) Festuca vaginalis subsp. cayambae (56b) Festuca vaginalis subsp. vaginalis (56a) Festuca venezuelana (16) Festuca woodii (11) 175 176 Festuca in South American Paramos ALPHABETICAL INDEX TO COLLECTORS Number in parentheses corresponds to number of the species in the taxonomic treatment as indicated in the Alphabetical List of Accepted Species and Subspecies preceeding this section. Acosta-Solis 21026, 20054 (4); 16963, 16974, 18839 (6); 110206,19821,19832,19835 (12); 6268 (13); 7581 (15); 9879, 18836 (19); 16358 (20); 9884, 9877(21); 18729 (25); 21031 (27); 16678,19006, 21439A, B (29); 5097, 19199 (31); 16765 (34a); 7554, 7614, 7623, 10538, 17670, 19196, 21195, 21208, 21216, 166989 (52); 21187 (56a) Aguirre & Merino 4219 (21) Aguirreetal. 4206(31) Ahumada, O. & A. Schinini 4085 (18) Alberto et al. 144 (6) Alfaro 1027 (la) Alfaro 1029, 1771 (54a) AlushShilomTon957(la) Anderson 1271 (14) Andre 3919 (29) Anonymous (20) Apollinaire & Arthur 18 (11); 9, 121 (24) Archer 3402 (6) Arechavalet s.n. (18) Argiiello 348, 354, 357 (52) Aristeguieta & Medina 3531, 3558 (54a) Aristeguieta2596(17) Aristeguieta 2603 (17) Aristeguieta 763 (8) Aristeguieta 947, 964, 1003, 2464, 2602, s.n. (54a) Asplund, E. 6359, 7156, 7535, 7559 (4); 6155, 6215, 7195, 7462, 7537, 7878, 16094, 16163, 17846 (6); 8150, 8425 (12); 7524, 8427 (19); 6020, 6400,6478,7558,7892,8447, 8470, 8606,16259, 17313,20217 (21); 6139,6151,6154,7392, 8135, 10375, 16161, 20176 (29); 7806, 7927, 8576, 8586, 10000, 17174 (31); 8446 (34); 7892, 8411, 8446, 8641 (34a); 8397, 8396 (39); 7926, 8594, 17334,17504 (41); 6134,6138,6140,6369,6370, 6349, 6348, 6471, 7055, 7803, 7880, 8191, 8402, 8639, 8645, 8644, 8647, 8718, 8724, 9634, 9964, 16807, 9967, 17022, 17176, 17522, 18286 (52); 3648,6620,7397,7497,7872,8399,8721,17338 (56a); 8648 (56b) Badilloetal. 7562(17) Baldillo 3496 (2) Balslev & T. de Vries 3471 (29) Balslev 1615 (31); 23600 (52); Balslev et al. 3901 (31); 3435, 3438, 3437, 3675, 3733, 3734, 3738, 4070 (52) Barclay & Juajibioy 7017, 7033 (lb); 10180 (17); 7961 (21); 6109, 6517 (24); 8178 (29); 6456, 8804 (31); 8905 (34a); 6313 (41); 8763 (45); 5942, 5945, 6052, 8085, 8162, 8264, 8325, 8641, 8819, 9142, 9160 (52); 9656, 9895, 10214 (54a); 8166, 8905 (56a) Barclay 6431 (22); 4529, 5865 (52) Barclay etal. 9677(17) Barford & Blicher-Mathiesen 41546 (52) Barreto 1871 (8); 656 (17); 577 (18); 645 (54a) Baruch46(17) Beaman, J. 2284, 2363, 3216, 3482 (54a) Becker etal. 1234(29) Bejarano44(36);22(38) Bell 17 (4) Benavides 6497 (19); 2660, 6554 (52) Benoist 2515, 4636 (52) Berg & Steinmetz 187 (54a) Berg 470 (17); 98-29-16 (54a) Bernal & Jimenez 1127 (19) Bernardi 268 (17); 18618 (18); 275 (54a) Berro5994(18) Black 46-605 (24); 32 (52); 278, 279 (56a) Boechat 41059 (18) Bono 5662 (17) Bourgeau, E. 1032 (la) Brade & Kuhlman 15631 (18) Brandbyge 42597 (34a); 42133, 42586 (52) Bravo 56, 58(21) Breedlove, D. 11151, 39265 (la); 26723 (54a) Brescia & Marches 4212 (8) Briceno2854, 3111,3119,3178, 3357, 3711 (54a) Briceno&Adamo 184,187,949,1067,1131,1139,1988 (54a); 2052 (54c) Briceno & Molinillo 3447 (54a) Briceno et al. 440, 1146, 2330, 2667, 3054, 3077 (54a) Briceno, B. & G. Adamo 2003 (2); 191, 196, 274, 306, 948, 1086, 1114, 1320, 1705 (3); 2011 (8); 286 (12); 244 (17) Briceno, B. & Molinillo 3461 (17) Briceno, B. 3074(17) Briceno, B. et al. 2328 (2); 2601, 3345, 3390 (3); 358 (17) Buddie s.n (13) Buendia 2 (29) Burger & Gomez 8246 (la) Burgeretal. 7428 (la) Burkart 19654 (18) Burman 880, 905 (8) Cabrera 438 (31) Camargo 59992 (8) Festuca in South American Paramos Camp 3327 (9) Cano 3780 (29) Carbono 2474 (lb) Castelano & Monasteries 20 (54a) Castellafios 24729 (8) Castellanos et al. 3 (24) Cazalet & Pennington 5417 (6); 5763 (31); 5749,5749B (41); 5434 (52) Ceron & Alarcon 4805, 12260 (52) Ceron & Montesdoca 12010 (52) Ceron 10605,15102,21862,25977(29); 12609, 18088, 28250, 28957, 29480 (52) Ceron et al. 11813, 19819,25697(52); 19822 (56a) Cerratel540(42) Chardon5010(31) Chase, A. 8276, 9659, 9672, 9673, 10101 (8); 8302 (18) Clark etal. 1372(6) Clausen, J. 34 (54a) Clayton4491(8);4301 (18) Cleef&Urribe6694(24) Cleef4376,8285(19); 181,1587,6258(24); 1048,1272, 3605, 5259, 7704 (28); 7139 (30); 9078 (37); 982A, 1090, 6285 (38); 7930 (53) Cleefetal. 5983(31) Cleef, A.M. 181 (24); 6826 (36) Clements etal. 2170(31) Correaetal. 129(22) Cuatrecasas, J. & Castaneda 24736 (lb); 24532, 24563 (51); 25136, 25133 (54b) Cuatrecasas, J. & Garcia-Barriga 10037, 10207 (29) Cuatrecasas, J. & H.G. Barriga 9970 (10) Cuatrecasas, J. & Jaramillo-Mejia 25742 (24); 25865 (28) Cuatrecasas, J. & Willard 26292 (52) Cuatrecasas, J. 9292, 23137, 23178 (22); 9311 (31); 7966, 9475, 10446 (38); 23201 (41); 14694, 19059 (52) Cuatrecasas, J., Espinal & Ramos 27562 (22); 3550 (24) D'Orbygny39(18) Davalos 22 (6); 19 (31); 18 (45); 21, 28 (52) Davidse, G. & D'Arcy 10244 (la); 11166 (8) Davidse, G. et al. 3223 (3); 11074, 11082, 11087 (8); 5545 (38); 19610 (55) Davidson 1043 (la) Degen861 (18) Diaz-Piedrahita & Jaramillo-Mejia 1821 (31); 1821(41) Diaz-Piedrahita & Rangel 2037 (22) Diaz-Piedrahita 2367 (24) Dillon, M. etal. 6359A(29) Dombrowski 2432 (8) Dusen 3624, 3515, 7783, 7808, 17561 (8); 8972 (18) Dutra 331, 402 (8); 538, 543 (18) Dziekanowski & Bolingbroke 1911 (54a) 177 Echeverry 1973 (22) Edwards s.n. (45); 70 (52) Ehrenburg 60 (21); 1, 27, 36, 191 (52) Ellenberg 7024 (7) Escalona & Gallegos 287, E384 (29) Espinal 81, 1035 (52) Etter662(29);661(35) Fagerlind & Wibom 934-bis (21); 934bis, 932 (56a) Fagerlind et al. 934bis (34a) Fegan & Falconi 9 (21) Fernandez-Perez 2931 (52) Ferreyra, R. 7526, 7281, 7642, 7664 (14); 8566 (42); 15512, 15518(52) Fierrp24(31) Figueredo 119, 126(19) Firminll6(29); 137(52) Forero, E. et al. 45, 82, 92 (12) Forrero etal. 3644(31) Fosberg, R. 20749 (19); 20745 (28) Fraser s.n. (4) Funck&Schlim 1132 (54a) Galeotti, H. 5766, 5778 (la) Gamboetal. 819 (la) Garcia-Barriga 7838A(12); 1016, 1019 (24); 1018 (28) Garzon617(38) Gentry 12378, 12685 (52) Goergem & Wagner 50197 (8) Gonzalez et al. 137,206 (54a) Gonzalez-Ledesma et al. 484,478 (la) Gregory & Eiten 301 (54a) Grignon 84189 (52) Grubbetal. 639 (31); 656 (52) Guerrero 152 (38) HalloyB-54(21) HammeLB. 5619 (la) Hanselmann & Loleveless 296 (54a) Harling & Andersson 12538 (52) Harling 11255 (52) Harling et al. 8904, 10415 (6); 10533 (21); 8635 (27); 8924, 10477 (52) Harlington 1197 (21); 4067, 4552 (52) Harteman 39 (6); 67 (52) Hartman7a(19) Hartweg 1455 (52) Hatschbach, G. 1912, 4715, 13820 (8); 9551, 20379, 20381,35575(18) Heller 306 (54); 62, 306 (54a) Hernandez 1022A(30); 1311 (36) Hernandez et al. 1336 (54a) Herrera 3730 (8) Hitchcock 28 (18); 496 (54a) Hitchcock, A.S. 21042 (4); 20798,21094,21512,21682 (6); 20746 (9); 21981 (21); 21926 (29); 21978 178 Festuca in South American Paramos (31); 21058 (41); 21944, 21951, 22030 (52); 22328 (42); 7165 (54a); 22019 (56a) Holm-Nielsen & Balslev 23755 (21) Holm-Nielsen & Jaramillo 28415 (52) Holm-Nielsen 1482, 24604 (6); 6612 (31); 24871 (34a); 1485, 5383, 5427, 6462, 20789, 25163 (52) Humbert 26353, 26506, 26806, 26824 (54a) Humboldt & Bonpland 2296, s.n. (9); s.n. (15); s.n. (19); s.n. (25); s.n. (29) Idrobo et al. 3026 (6); 3678 (19); 3682 (21); 3737 (29); 3834 (52) Infantes 435 (14) Izuriete 129 (29) J.F. Smith 1997, 1999(52) Jahnl8,43(54a) Jahn,A. 62(17) Jameson 14, 296 (21); 191 (29); 230 (41); 70, 91, s.n (52); 230 (56a) Jaramillo - Mejia & Cleef 5738 (22) Jaramillo & Zak 623 (52) Jaramillo et al. 11855, 11856(27) Jaramillo-Mejia 6237 (31) Jaramillo-Mejia et al. 5634, 5683, 5770 (31); 5747, 6149,6210(41) Jorgensen, P.M. 2074 (52) Jorgensen, P.M. et al. 2099 (52) JRJ895(la) Juan &Alva 101 (54a) Jurgens 6258 (8); 175(18) JurgensenG85(18) K.T. Hartweg 1450 (56) Killip & Smith 17861 (6) Killip 6709 (6); 19719 (16); 17329 (24); 18479 (26); 18479(35) Killip et al. 38050 (38) Kjell von Sneidern 362, 435, 415, 2144 (52) Klein 3872 (18) Kozera & Izernhagen 476 (8) Kuhlmann 2242 (8) L. Parodi 5673 (8) L.G. Clark & X. Londono 376 (41) Laegaard & Aguirre 20611 (43) Laegaard & Sklenaf 20308, 20287 (15) Laegaard, S. & Aguirre 20609 (21); 20608 (49b) Laegaard, S. & Grignon 19395 (52) Laegaard, S. & Mayorga 17536(38) Laegaard, S. & S.A. Renvoize 70580, 70577 (6); 70519 (21); 70485, 70542, 70873, 70882 (29); 70511 (31); 70555 (41); 70558 (56a); 70488 (56b) Laegaard, S. & Sanchez 20027 (21); 20033 (29) Laegaard, S. & Sklenaf 20302 (9); 20301 (12); 20335, 20363 (21); 20296 (29); 20282A (52) Laegaard, S. 54540A, 54540B, 102316, 54164, 55260, 55281 (4); 18748, 18727, 18483, 20688, 52285, 52754, 53118, 53372, 55282, 101406, 101711, 101781, 102227, 102293, 102318, 105069, 105132 (6); 20405, 102535, 105256, 105235 (9); 18623, 18667, 18704, 18744, 101818, 102716 (12); 55290, 55411, 55567 (15); 5866, 54327, 54892, 101731 (19); 18736, 19739, 52374A, 52605, 52607, 53580, 53837, 54148, 55301, 55535, 55673, 71737, 101384, 101432, 101718, 102087, 102268, 102237, 103107 (21); 101260 (25); 53965, 53597 (27); 20379, 52301, 54802, 54809,5556670002,70014,70016,71031,71695, 55410,101131,101350,101355,101340,101814, 101856, 102164, 102285, 102857, 105102 (29); 18449, 51318, 53289, 53343, 54581, 54972, 55425, 55680, 101284, 101383, 101595, 101695 (31); 55769,101162,101294,101716(33); 19155, 52125, 54594, 54820, 55346, 55736, 101583, 102156, 102892 (34a); 101517, 101612, 102253 (34b); 53293, 53307, 53869, 54518B, 55696, 55718, 70491, 102797, 102803, 102889 (41); 19452, 54548, 55368, 55698, 101194, 101373, 101692, 102262, 102666, 102871, 103105 (45); 70309 (48); 19148, 19395, 21016, 51060, 52107, 52373, 52837, 53125, 53213, 53373, 53834, 53871, 54351, 54409, 54416, 54506, 54731, 55001, 55276, 55279, 55286, 55304, 55401, 55417, 55452, 55635, 55672, 55734, 55748, 55771, 69261, 71739, 71741, 101181, 101164B, 101264A, 101577, 101726B, 101856, 102103, 102100, 102134B, 102376 (52); 19159, 19598, 51386, 52202, 52656, 52659, 55327, 55400, 55536, 55735, 101335A, 101460, 101495, 101559, 102096, 102149, 102743, 102758, 102795, 102796, 102848, 103127 (56a); 19589, 101493, 101505, 101828, 105123 (56b) Laegaard, S.etal. 18855,19095,70813,103010,103079, 103032 (6); 103003 (12); 20994 (20); 19066, 20824, 20993, 102737 (21); 18672 (29); 18606, 18859,19280,20733,20931 (31); 102744,102739 (34b); 19630, 102760, 102725 (41); 53859 (45); 19285 (49b); 18868,20613,102733,103067(52); 70553, 70554 (56a) Langsdorff s.n. (8) Larsenet al. 21 (31) Leavenworth & Hoogstraal 1218 (54a) Leavenworth et al. 1213 (la) Lechler2114(29) Leite3490(18) Leiva & Leiva 96 (14); 1094 (42); 2525 (52) Leiva 797 (14) Lenll39(31) Leon, B. & Young 1662 (31); 1101, 1364,1679 (52) Liebmann6109,6110,12904(la); 517,6108 (54); 2886, 3030, 6117 (54a) Festuca in South American Paramos Linderman et al. 4658 (8) Lindig 1116, 1117 (6); 1862 (8); 14 (19); 1013, 1053 (24) Littele 15712 (54a) Little 15717 (17) Liana 71 (31) Longhi & Sampaio 368 (18) Longhi&Witten5011 (18) Lopez & Sagasteguii 3340, 8180 (31); 2827 (42); 8180 (52) Luteyn, J. & Luteyn 5713 (52) Luteyn, J. 13452 (52) Luteyn, J. et al. 13051 (22); 12834, 12842 (52) Lyonnet 260, 2973 (54a) MacDougal et al. 4504 (6) Maf57(52) Maguire 3487 (8) Manrique, E. et al 1147 (la) Martin 630 (la) Matuda 25909 (la); 2337, 2360 (54a) Meier 3025 (17) Meier et al. 3026 (54a) Mejiaetal. 6181 (22) Mexia 2684 (54a) Mille s.n. (4); s.n. (15); 664, 659 (25); 283, 285 (29); s.n. (52) Millesetal. 281(6) Molau & Eriksen 3244 (31); 3298 (41) Montesdeoca 280, 282, 284, 295, 339, 348, 585 (52) Morales 5194 (54a) Mora-Osejo 3195, 3503(52) Morrone, O. & A.M. Cialdella 854 (8) Mueller 2115 (la) Munoz 207 (52) Mutis 5544 (6); 5570 (24) Naranyo4(31) Narvazez et al. 548(31) Nee, M. & Soule 33022 (54a) Neill, D.A. et al. 11992, 12042 (31) 011gaard, B. & Balslev 10074, 8170, 8545, 8868, 9924, 10122 (31); 9921, 10073,10100 (45); 8926 (52) 011gaard, B. & Holm- Nielsen 38728 (31) 011gaard, B. et al. 34039 (27); 34202, 38540, 38541, 38670(31) Ortiz 1272 (52) Oxford Expedition 30 (21) Palicel,3,4,9(6);6,7, 8(52) Pedersen, T.M. 819, 11475 (18) Pedraza et al. 491 (24); 228, 242, 278 (28) Penafiel 1067 (52) Penafiel et al. 69 (29); 402, 527, 667, 720 (52) Penland & Summers 583 (21); 889, 969 (52) 179 Pennell 9951 (21); 3003 (22); 1923, 2018 (24); 13231 (29); 3064, 9890 (31); 13552,13845 (52) Pennell, F. 7057 (52) Peterson, P.M. 11181 (12); 8815 (29); 8721 (52) Peterson, P.M. 14224 & O. Tovar (29) Peterson, P.M. 14907 &NRefulio Rodriguez (21); 14892 & N. Refulio Rodriguez (31); 13875, 13961, 14011,14936,14855,14925 &N. Refulio Rodriguez (42); 13951, 14901 &N. Refulio Rodriguez (43); 13922,14953,14955,14957,14968,14989, 14992, 15178 & N. Refulio Rodriguez (29) Peterson, P.M. 8872, C.R. Annable & M.E. Poston (6); 8858, 8870, C.R. Annable & M.E. Poston (21); 8717, 8734, 8747, 8846, C.R. Annable & M.E. Poston (29); 8768, C.R. Annable & M.E. Poston (34a); 8744, 8759,8770,8846,8865,8876,8932, C.R. Annable & M.E. Poston (52); 8739, C.R. Annable & M.E. Poston (56a) Peterson, P.M. 8973, 8996 & C.R. Annable (21); 8979, 8995, 8996A & C.R. Annable (34a); 8982, 8973 & C.R. Annable (34b); 8951, 8983, 8985 & C.R. Annable (52); 8987 & C.R. Annable (56a); 8984, 9000 & C.R. Annable (56b). Peterson, P.M. 9118,9139, E.J. Judziewicz & R.M. King (21); 9076, 9154, E.J. Judziewicz & R.M. King (31); 9086, E.J. Judziewicz & R.M. King (41); 9122, 9127, 9140, 9155, E.J. Judziewicz & R.M. King (52) Peterson, P.M. 9209,9235, E.J. Judziewicz, R.M. King & P.M. Jorgensen (21); 9179, E.J. Judziewicz, R.M. King & P.M. Jorgensen (29); 9204, E.J. Judziewicz, R.M. King & P.M. Jorgensen (31); 9197, E.J. Judziewicz, R.M. King & P.M. Jorgensen (45); 9189, 9215, 9245, E.J. Judziewicz, R.M. King & P.M. Jorgensen (52); 9228, E.J. Judziewicz, R.M. King & P.M. Jorgensen (56a) Peterson, P.M. 9313 & E.J. Judziewicz (6); 9259 & E.J. Judziewicz (9); 9297, 9316, 9512 & E.J. Judziewicz (29) Pinto-Escobar & Hernandez 523 (24) Pinto-Escobar 415, 1845 (31) Pittier 1511 (6); 437, 1320, 1444 (9); 13276 (17); 1409 (52) Ponce & Trujuillo 245 (12) Prescott 913 (6); 656, 834, 839, 848 (52) Pringle, CG. 3945 (la) Puiggari s.n. (8) Quarin3409(18) Quintana 2 (52) Quipuscoa & Vilchez 2611 (52) Quiroz 1929 (29) Rambo 36446, 49315, 54011, 54999, 54989 (8); 45452, 45466, 53471, 53816, 54704, 60082 (18) Ramirez 1404 (9) 180 Festuca in South American Paramos Ramirez, M. & N. Cuello 3523, 3524, 3528 (5) Ramsay & Smith 245,247 (21); 415B, 502,785 (31); 410 (41); 343 (45); 422, 510, 742, 1016 (52) Ramsay 896 (31); 1035 (45); 426 (52) Ramsay et al. 104A(21); 153 (45); 1057 (52) Rangel & Aguirre 3720 (24) Rangel 1650(28) Rangel et al. 945 (lb); 1800 (31); 3515 (36); 1732, 1791 (41) Rauh & Hirsch E321, E331 (52); E315 (56a); RegnellHI 1409, s.n. (18) Reitz & Klein 10832, 10399, 12509, 14611, 14585 (8); 11812, 11948, 14172(18) Reitz 2611 (8) Renvoize, SA. & S. Laegaard 5031 (50) Renvoize, SA. 4879 (29) Ricardi & Carrez 5930 (17) Rimbach54(15);97a(56a) Robinson & Beltran 3142 (19) Rodriguez 4130 (38) Roldan et al. 360 (6); 396, 307 (19) RosengurttB-2764(18) Ross 1276 (54a) Ruiz & Romero 3 (30) Sagasteguii 14785 (7); 2991 (14) Sahn&Hernanl060(19) Saint-Hilaire312,304(8) Salamanca AC 172 (31) Salamanca et al. 40, SS143 (41) Sampaio & Arzivenco 341, 362 & 404 (18) Sanchez & Hernandez 950 (19) Sanchez 283 (24) Saravia3912(38) Schinini, A. & Carnevali 10523, 10687 (18) Schulz & Rodriguez 317 (54a) Schulz317(54a) Schwaike s.n. (18) Scito&Kuniyoshil315(8) Scur 656 (8) Seaton 228 (54a) Seifriz 489 (lb) Sellow s.n. (8) Sellow.F. s.n. (18) Sklenaf & Cruz 8706 (21) Sklenaf & Kosteckova 1-24, 15-5, 16-1, 66-10, 85-8, 86-9, 87-6,1905 (21); 64-19,91-8,97-10,112-25, 1028, 1106, 1142, 1478, 1817, 1890 (31); 1-24, 858,1322 (34a); 2738 (39); 49-19,55-2,56-2,9-1, 735, 1836, 1853, 1863, 1893, 1903, 1931, 1933 (41); 956,1031,1929,2168(45); 14-9,1881 (52); 143-2, 198 (56a) Sklenaf & S. Laegaard 7092 (52) Sklenaf & Sklenafova 2168, 2262 (21); 2990 (41) Sklenaf & Zapata Cruz 8705 (43); 8630 (52) Sklenaf 2596, 8204 (45); 8067, 8115 (56a) Smith & Klein 10887, 10956, 11384, 11335, 11611, 11836, 12024, 12188 (8); 10855, 10958, 13674, 15474, 15761, 15805, 15916, 15936(18) Smith & Reitz 10127, 10428 (18) Smith 10561 (7); 7534, 8233, 10537, 10769, 11389, 12110 (14); 6222 (29); 8284, 11318 (31); 9942, 11493 (42); 3482 (52) Smith et al. 9389 (18); 7314, 7321a, 7332 (21); 3627 (29); 7428, 8879, 9225, 11127 (31); 3312, 4278, 9939, 9944, 12630 (42); 11851 (43); 3278, 9783, 10229, 10253, 11799(52) Sneider 430 (6) Soderstrom, T.R. 1229 (24); 1307 (28) Sodiro 36/11, s.n. (4); 36/6, 282, s.n. (6); s.n. (9); s.n. (12); s.n. (13); 20/5, s.n. (15); s.n. (25); s.n. (29); s.n. (41); s.n (52); s.n. (56a) Sodiro, A. 36/4(41) Sohn&Matuda991(54a) Sosa et al. 43 (29) Sparre 13562 (4); 15076, 16950, 16920 (6); 15635, 15708, 15871 (21); 15827, 15836, 16006 (29); 17719 (31); 13709, 15781, 15828, 17408 (52) Spruce 5938 (6); 5509 (52) Stancik, D. & Carvajal 1866 (23); 1783, 1787, 1811, 1848, 1874, 1891 (24) Stancik, D. & Galvis 2025, 2026, 2028 (46) Stancik, D. & Medina 2577 (23); 2166 (32) Stancik, D. 4173, 4174, 4177B, 4180, 4281 (2); 4171, 4175, 4178, 4215, 4250, 4256, 4257 (3); 3668, 4104 (4); 4286 (5); 2632,2754,2772,2776,2858, 2866,2870,2902,2992,2991,3010,3292,3352, 3595, 3667, 3656,3771, 3781, 3792, 3885, 3886, 3985,4095,4105,4113 (6); 4177,4179 (8); 3221, 3309,3310,3798,4102 (12); 3317(15); 4262 (16); 4182, 4192, 4248 (17); 948, 1455, 1365, 2024, 2190, 2775,2983, 3412, 3399, 3554, 3551, 3685, 3683,3705,3712,4092, s.n. (19); 3457 (20); 2744, 2981,3004,3108, 3128,3320,3358,3359,3373, 3380,3416,3455, 3596,3610,3686,3713,3787, 3780,3800,3802, 3808,3858,3879,3887,3903, 4023,4028,4031,4090B, 4093,4094,4111,4158, 4162 (21); 3375, 3376, 3377, 3404, 3405, 3599 (22); 303, 928, 1065, 1069, 1287, 1288, 1289, 1290, 1297,1302, 1319, 1321,1323, 1324,1325, 1352, 1355,1357, 1391, 1402,1436, 1512,1516, 1530, 1531,1546, 1842, 1851,1854, 1921,1923, 1924,1969,1978,2003,2025,2052,2053,2063, 2073, 2168,2174, 2192, 2218,2256, 2408,2487, 2507, 2508, 2511, 2512, 2527, 2550, 2560,2561, 2562, 2563,2564, 2565, 3480, 3481, 3491, 3492, 3493,3494,3496, 3500,3538,3540,3542,3544, 3545, 3546, 3560 (24);4290, 4289 (26); 3365, 3983 (27); 210,217,218,224,225,226,228,231, 241, 248,250, 256, 257, 258, 264, 265, 267, 268, 270, 277, 278, 302, 311, 312, 313, 314, 315, 316, 317, 318, 319, 320, 321, 491, 3494, 3559, 3561, Festuca in South American Paramos 3562,3563,3564,3565,3569,3568,3579,3580, 3583,3584,3585,3586,3587,3589,3590,3566, 3567, 3570, 3572, 3573, 3574, 3575, 3576, 3577, 3578, 3581, 3582, 3585, 3588 (28); 1788, 3312, 3313, 3314, 3317, 3793, 3794, 4106 (29); 2724, 2759, 3044, 3050, 3241, 3279, 3401, 3402, 3410, 3411, 3606, 3611,3612,3613,3614,3637,3675, 3703,3763,3767,3849,3871,3904,4097,4150, 4151 (31); 1493 (32); 948,1365,1404,1418,1533, 2371 (30); 3080, 3710, 3711, 3719, 3720, 3880, 3883,4024,403 IB, 4033B, 4034 (34a); 272,409, 466, 467, 924, 926, 945, 927, 1339, 1366, 1373, 1407,1421, 1422, 1423,1438, 1457,1463,1465, 1482,1503, 1532, 1555,1556, 1576,1577,1580, 1628,1602,1635,1640,1629,1688,1689,1690, 1827,1865,1828,1830,1920,1935,1950,1964, 2000,2020, 2045, 2051,2050, 2054,2130, 2136, 2137,2138, 2160, 2188,2189, 2224,2230, 2232, 2236,2240, 2495, 2502,2503, 2505,2249, 2257, 2313,2332, 2314, 2346,2350, 2367,2372, 2373, 2489 (36); 1886 (37); 1286, 1347, 1360, 1363, 1396,1402,1511,1513,1523,1922,1929,1939, 1951,1952, 1944, 1945,1985, 1986,1987, 1988, 2005, 2019, 2109, 2115, 2116, 2117, 2120, 2114, 2136, 2250, 2471, 24732472, 2597, 2599, 2612, 3539 (38); 4287, 4288 (40); 3031, 3034, 3048, 3373B, 3374,3378,3383,3392,3393,3394,3413, 3601, 3602, 3607,4039,4122,4129,4133,4134, 4166,4149 (41); 2056,2749,2783,2786,3054A, 3054B, 3098,3101,3228,3235,3267,3898,3899, 3990,4081,4090,4091,4092B, 4098 (45); 340 IB (47); 3846, 3859, 3873 (49a); 2613, 2618, 2617, 2619,2631, 2656, 2657,2658, 2659,2685, 2748, 2764,2766, 2789, 2790,2802, 2856,2876, 2860, 2962,2963,2908,2909,2910,2911,2918,2914, 2917, 3013, 3119, 3146, 3147, 3149, 3150, 3151, 3152,3153,3155,3158,3162,3164,3169,3189, 3190,3193,3176,3240,3326,3327,3335,3336, 3337,3344,3351,3353,3354,3355,3360,3363, 3367,3608,3609,3671,3677,3687,3688,3690, 3691,3692,3704,3705B, 3707,3708,3709,3714, 3715, 3716, 3717, 3769, 3770, 3772, 3773, 3774, 3804,3805,3811,3815,3837,3847, 3848,3850, 3862,3863,3864,3865,3872,3882,3884,3888, 3889,3900,3901,3902,4026,4027,4030,4032, 4089, 4096, 4101, 4103, 4115, 4163, 4164 (52); 4172,4176,4181,4191,4207,4225,4242,4243, 4279, 4280 (54a); 4258, 4259 (54c); 3001, 3014, 3020,3134,3137,3323,3341,3706,3718,3881, 4013 (56a); 3182, 4033, 4035 (56b) Standley 61885, 61899, 65227 (la) Steyermark, J. & Koyama 102389 (17) Steyermark, J. 53481 (31); 53146, 53182 (52) Steyermark, J. A. 53092 (49a) Storek 2889 (54a) Stiibel 297 (29); 257 (52) 181 Stiibel, A. 297 (29); 207 (52) Sturm 33(38) Swallen, J.R. 8033, 8168, 8231, 8540, 8859 (8); 8089, 8189(18) Tamayo2326(16) Taylor et al. 170 (54a) Tenorio 15093 (1) Teran & Melchior 833, 848 (17) Teran 7732, 7909, 13215(17) Tillet & Honig 746-765 (lb) Tillett, S.S. 747-1122 (54b) Torres-Romero & Lozano 2982 (28) Torres-Romero et al. 1875 (31) Tovar Serpa, O. 2057 (7) Tovar, O. 143 (14) Triana810, s.n. (31);s.n. (38) Tsugaru & Otsuka B-2265 (18) Turpe, A.M. 2932(8) Tutin 1285 (29) Ule 243, 616, 1959(18) Valderrano 34 (38) Vails 56, 2386 (8) Vails etal. 1893(18) Vareschi 1232 (17); 2134, 2232, 2244, 2248, 3853, 7021 (54a) Vareschi et al. 1233(17) Vargas 11176 (52) Vargas et al. 2083, 2583 (52) Vega & Ruiz 3584 (29) Vega 249 (7); 1966 (14); 5419, 7143 (21); 723, 2250, 2576, 3239, 3830, 4661, 5013 (29); 7087 (31); 2263, 4168, 4371, 5271, 5427, 6616, 10509, 10764, 10889 (42); 4378, 7111, 7124 (43); 1385, 1969,4043,4145,7136,11181, 11190(52) Vega etal. 1544(7); 1638, 10320 (21); 411, 1393,1598, 3283, 3855, 5185, 5696, 5745, 6933 (29); 1695, 2012, 2061, 2527, 3797, 5744, 6694, 6974, 6975 (42); 2018, 3494, 6914, 6955, 6994, 7041, 7128 (43); 7011 (50); 2011,2464, 7062, 10795, 10904, 10906(52) Velez etal. 4514(22) Vivar & Marin 1305 (12); 1306 (13) Vries, T de s.n. (45); s.n. (52) Wagner 944, 946, 947, 5010 (8) Wagner et al. 298 (8) Wawra 947 (54a) Weberbauer, A. 3230 (14); 3275 (42); 3974 (52) Whymper 1326, s.n. (31); 1330 (41); 1322, 1638 (52) Widgrens.n. (18) Wingeetal. 1272, 1362(8) Wood4326,4347,4792,5066,4648 (6); 3842, 5104 (8); 5309 (9); 4639 (12); 5259 (17); 4247, 5331 (21); 182 Festuca in South American Paramos 4033 (22); 3612 (24); 4539 (28); 4797 (29); 4481, 4507 (30); 4030,4062,4063,4802 (31); 5141 (36); 5260 (37); 3523, 4539 (38); 4806, 4807 (52) Wood, J.R.I. 5254, 5354 (11); Wurdack, J. 1572 (31); 1571 (52) X. Londono et al. 550 (23) Yepes-Agredo 760 (22) Young & Watson 3484 (52) Zapata 16506, 16508 (43) Zardinii 7701 (18) Zuluoga 107 (38) Festuca in South American Paramos 183 INDEX TO SCIENTIFIC NAMES Accepted names are in regular type; main entry for each is in bold face; synonyms are in italics; figures are in parentheses. Avena secunda Salisb. 42 Bromus arundinaceus (Schreb.) Roth 42 caldasii Kunth 36 depauperatus J. Presl 5 elatior (L.) Koeler 42 littoreus Retz. 42 pratensis (Huds.) Spreng. 44 procerus Kunth 36 Bucetum elatius (L.) Parnell 42 pratense (Huds.) Pamell 44 Diplachne procera (Kunth) Spreng. 77 Festuca L. 11 aequipaleata E. Fourn. 136 americana (Pers.) F. G. Dietr. 44 ampliflora Doll 52 amplissima Rupr. 5, 6, 13, 16, 18, (19, 20, 147) amplissima Rupr. subsp. amplissima 17 amplissima subsp. magdalenaensis 6 amplissima subsp. sierrae Stancik 18 andicola Kunth 5, 6, 7, 12, 16, 22, (54), 55, (56, 158) arundinacea Schreb. 12, 16, 42, (43, 45, 153, 154) arundinacea subsp. orientalis (Hack.) Tzvelev 42 asplundii E.B. Alexeev 7, 14, 17, (83), 84, (85), 104, 136,(163) Festuca sect. Aulaxyper Dumort 16, 83, 106 azucarica E.B. Alexeev 13, 16, 63, (64, 65, 159) boyacensis Stancik 6,15, 17, (79), 88, (89), 91, (164) breviaristata Pilg. 6, 129 breviglumis Swallen 16, 38 cajamarcae Pilg. 129, 130 caldasii (Kunth) Kunth 5, 7, 12, 16, (35), 36, (37), 38, 40,(151,152) carchiense Stancik 6, 15, 17, 88, (90), 91, (93), 108, 122, (164) casapaltiensis Ball 7 Festuca sect. Cataphyllophorae E.B. Alexeev 6, 16, 65, 70, 77, 83 chimborazensis E.B. Alexeev 15, 17, 88, 91, (92, 93, 164, 165) chimborazensis subsp. chimborazensis (92), 93, 94 chimborazensis subsp. micacochensis 6, 93, 94 chita Stancik 6, 13, 17, 95, (96,102) chitagana Stancik 6, 12, 16, (61), 66, (67) cleefiana E.B. Alexeev 14, 17, 75, (93), 95, (97, 165) cocuyana Stancik 6, 15, 17, (83), 98, (99), 100, 126, (166) colombiana E.B. Alexeev 13, 16, (65), 66, (68), 69, 72,(159,160) coromotensis B. Briceno 5, 13, 16, 20, 21, (20, 21), 22, 28, (147, 148) cundinamarcae E.B. Alexeev 14, 17, 100, (101, 102, 166, 167) dasyantha Kunth 5, 7, 13, 16, (65), 70, (71), 72, 75, (160) densipaniculata E.B. Alexeev 14, 17,102, (103), 104, (106, 167) dichoclada Pilg. 6, 7, 12, 16, 45, (45), 46 dinirica Stancik 6, 7, 14, 17, (93), 104, (105), 106 dissitiflora Steud. 55 distichovaginata var. cajamarcae (Pilg.) St.-Yves 129 dolichophylla J.Presl. 6, 7 Festuca subg. Drymanthele V.I. Krecz. & Bobrov 16 drymeja Mert. & W.D.J. Koch 16 elatior L. 42 elatior subsp. arundinacea (Schreb.) Celak. 42 elatior subsp. arundinacea (Schreb.) Hack. 42 elatior subsp. arundinacea var. genuine subvar. orientalis Hack 42 elatior subsp. pratensis (Huds.) Hack. 44 elatior var. arundinacea (Schreb.) Wimm 42 elatior var. pratensis (Huds.) A. Gray 44 elviae B. Briceno 5, 12, 16, 22, (23, 26), 55, (148) eminens Kunth 6, 36 Festuca subg. Erosiflorae E.B. Alexeev 16 ferreyrae Tovar 129, 130 fimbriataNees 12, 16, 52, (53, 54, 157) flacca Hack, ex E.B. Alexeev 12, 16, (20), 24, (25), 33, (149) flexuosa Willd. ex Kunth 46 fluitans var. pratensis (Huds.) Huds. 44 fragilis (Luces) B. Briceno 5, 7, 14, 16, 48, (50, 51), 51,(156) fratercula Rupr. ex E. Fourn. 17 glabra Spreng. 44 Festuca sect.Glabricarpae E.B. Alexeev 6, 16, 38 glumosa Hack, ex E.B. Alexeev 7, 16, 17, 91, (102), 106, (107), 108, 117, 122, 126, 136, (168) glyceriantha Pilg. 14, 17, (106), 109, (110), 111, (168) guaramacalana Stancik 6, 7, 13, 16, 26, (26, 27), 28, (149, 150) hatico Stancik 6, 12,16, (61), 72, (73, 160, 161) Festuca subg. Helleria E.B. Alexeev 6, 16, 51 holubii Stancik 6, 16, 17, (106), 111, (112), 113 huamachucensis Infantes 15, 17,113, (114, 115) imbaburensis Stancik 6, 15, 17, 91, 108, 115, (115, 116), 117, 122,(169) laegaardii Stancik 6, 11, 17, 65, (65), 72, (74), 75, (161) 184 Festuca in South American Paramos leptothrix Trin. ex Doll 33 liebmannii E. Fourn. 136 littoralis Wahlenb. 42 livida (Kunth) Willd. ex Spreng. 5, 16, 51 Festuca subg. Mallopetalon (Doll) E.B. Alexeev 6, 16 monguensis Stancik 6, 14, 17, (106, 118), 119, (169) multiculmis Steud. 136 nereidaensis Stancik 6, 14, 17, (115), 119, (120, 170) orgyalis Willd. ex E. Fourn. 77 orgyalis Willd. Ex Spreng. 77 orientalis (Hack.) Krecz. & Bobrov 42 oroana Stancik 6, 14, 17,119, (121), 122, (126, 170) orthophylla Pilg. 7 ovina L. 11 ovina subvar. jamesonii St.-Yves 106 parciflora subsp. loxana 6, 122,124 parciflora subsp. parciflora 122, (123), 124 parciflora Swallen 15, 17,122, (126) peruviana E.B. Alexeev 7, 77, pilar-franceii Stancik 6, 13, 17, 65, 69, 75, (76, 79, 162) pinetorum Swallen 5, 7 poaeoides Michx. 44 poaeoides var. americana Pers. 44 pratensis Huds. 12, 16, (40), 44 pratensis var. elatior (L.) Gaudin 42 presliana Hitchc. 5 procera Kunth 5, 6, 13, 16, 17, 72, 77, (78, 79), 79, 84,(162) procera Nees & Meyen 77 pseudosclerophylla Krivot. 42 quadridentata Kunth 5, 7, 12, 16, 46, (47, 51, 154, 155) quadridentata subsp. eminens (Kunth) St.-Yves 36 quadridentata var. caldasii (Kunth) St.-Yves 36 quadridentata var. dichoclada (Pilg.) St.-Ives 45 quitensis Willd. ex Kunth 36 racemosa Willd. ex Spreng. 55 reclinata Swallen 12, 16, 38, (39, 40, 152) regeliana Pavl. 42 renvoizei Stancik 6, 15, 17,124, (125, 126) rigescens (J. Presl) Kunth. 7, 55 rubra L. 13, 16, (51), 57, (58, 157, 158) Festuca sect. Ruprechtia E.B. Alexeev 6, 16, 18 sanctae-martae Stancik 6, 15, 17, 100, (115), 124, 126,(127, 171) scabra Willd. ex Steud. 77 Festuca sect. Schedonorus (P. Beauv.) Koch 16 Festuca sect. Schedonorus (P. Beauv.) Peterm. 11,16 sodiroana Hack, ex E.B. Alexeev 7, 12, 16, 21, 22, (26), 28, (29, 150) soukupii Stancik 6, 14, 16, 55, 59, (60, 61, 158) steinbachii 48 sublimis forma vivipara St.-Yves 146 sublimis Pilg. 77 subulata Trin. 16 subulata v&r.fraseriana St.-Yves 24 Festuca subg. Subulatae (Tzvelev) E.B. Alexeev 16 Festuca sect. Subulatae Tzvelev 16, 28 subulifolia Benth. 14, 17, 84, 104, 126, (128, 129), 130,139,(171) sumapana Stancik 6, 15, 17, 91, 122, (129), 136, (137) tenuiculmis Tovar 55 toca Stancik 6, 12, 17, 65, 81, (82, 83), 83, 84, (163) tolucensis Kunth 5, 14, 17, 84, 104, (129), 130, 136, 139,141,143 tolucensis subsp. culata Stancik & P.M. Peterson 6, 139,141 tolucensis subsp. perijae Stancik 6, 139, 141 tolucensis subsp. tolucensis (138), 139, (172) tolucensis var. subulifolia (Benth.) St.-Yves 126 tovariensis Stancik & P.M. Peterson 12, 16, 24, 31, (32, 35) turimiquirensis Stancik & P.M. Peterson 6, 14, 17, (102), 141, (142), 143 ulochaetaNees ex Steud. 6, 7,13,16, 24, 31,33, (34, 35,151) vaginalis (Benth.) Laegaard 7, 15, 17, (126), 143, (172) vaginalis subsp. cayambae Stancik 6, 143,146 vaginalis subsp. vaginalis 143, (144), 145 venezuelana Stancik 6, 13, 16, (45), 48, (49, 155) woodii Stancik 6, 13, 16, 36, 38, 40, (40, 41, 153) Gnomonia elatior (L.) Lunell 42 fragilis Luces 48 fragilis (Luces) Rauschert 48 Leucopoa pseudosclerophylla (Krivot.) Bor 42 Lolium arundinaceum (Schreb.) Darbysh. 42 pratense (Huds.) Darbysh. 44 Poa elatior (L.) Moench 42 hybrida var. vallesiaca Bronm. 42 phoenix Scop. 42 Schedonorus americanus (Pers.) Roem. & Schult. 44 arundinaceus (Schreb.) Dumort. 42 caldasii (Kunth) Roem. & Schult 36 elatior (L.) P. Beauv. 42 littoreus (Retz.) Tzvelev 42 phoenix (Scop.) J. Holub 42 pratensis (Huds.) Beauv. 44, 45 procerus (Kunth) Roem. & Schult. 36 radicans Dumort. 44 Tragus elatior (L.) Panz. ex B.D. Jacks. 42 pratensis (Huds.) Panz. ex B.D. Jacks. 44 Uniola effusa E. Fourn. 17 muelleri E. Fourn. 17 Vulpia ulochaeta Nees ex Doll 33