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 Contri- butions 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. turimiqui- rensis 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 Con- tributions 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 estu- diada, 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 discon- tinued 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 5 HISTORICAL OVERVIEW 5 TAXONOMY AND DIVERSITY 6 BlOGEOGRAPHY 6 MATERIAL AND METHODS 7 ACKNOWLEDGEMENTS 8 REFERENCES 8 TAXONOMIC TREATMENT 11 KEY TO THE SPECIES OF FESTUCA IN BRAZIL, COLOMBIA, ECUADOR, AND VENEZUELA 11 SURVEY OF THE SPECIES AND SUBGENERIC TAXA OF FESTUCA IN SOUTH AMERICAN PARAMOS 16 DESCRIPTIONS OF SPECIES OF FESTUCA FROM SOUTH AMERICAN PARAMOS 17 EXCLUDED NAME 146 SCANNING ELECTRON MICROGRAPHS OF LEAF EPIDERMAL SURFACES 147 APPENDICES 173 INDEX TO FIGURES 173 ALPHABETICAL INDEX OF ACCEPTED SPECIES AND SUBSPECIES 175 ALPHABETICAL INDEX TO COLLECTORS 176 INDEX TO SCIENTIFIC NAMES 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 first Venezuelan paramo flora by Vareschi (1970). is one of the largest genera in the Poaceae (Clayton Later, Alexeev (1986) in his work on Festuca for & Renvoize 1986; Watson & Dallwitz 1992). The Venezuela, Colombia, and Ecuador mentioned only genus has cosmopolitan distribution and it is an one species from Venezuela (F tolucensis). The important component of grass ecosystems of the recently published study of Briceno and Murillo temperate zone as well as the alpine grasslands (1994) treats Festuca in the state Merida (Ven- of the tropical zone. Although the diversity of the ezuela) and lists five native species (F. pinetorum genus Festuca is centered in the Holarctic zone of Swallen, F elviae Briceno, F tolucensis, Ffragilis, Eurasia and North America, with about 140 species F coromotensis Briceno). Stancik and Peterson (Darbyshire et al. 2003), the South American Andes (2003) indicated that Fpinetorum was misidentified represent a significant center of Festuca diversity. and therefore does not occur in Venezuela. In addi- tion, Davidse et al. (1994) mentioned the species HISTORICAL OVERVIEW.•In South America, taxo- F. amplissima Rupr.; the only known Venezuelan nomic studies of Festuca have been concentrated specimen (Tillett & Honig 746-765, MO) was in the southern part of the continent, i.e. Argentina, collected in Serrania de Perija on the Colombian- Bolivia, Chile, and Peru (Phillipi 1859-60, 1865; Venezuelan boundary. The earliest collections of Pilger 1898, 1906; Hitchcock 1927; Saint-Yves Festuca from Ecuador and Colombia were made by 1927; Parodi 1935,1953; Macbride 1936; Infantes A. von Humboldt and A. J. A. Bonpland (Expedition 1952; Foster 1966; Tiirpe 1969; Vareschi 1970; to Ecuador in 1802-1803; see Humboldt et al. 1816 Correa 1978; Matthei 1982; Tovar 1993; Brako and Jorgensen et al. 1999) and J.C. Mutis and F.J. & Zarucchi 1993; Zuloaga et al. 1994; Renvoize Caldas (Real Expedicion Botanica 1783-1808; see 1998; Soreng et al. 2003). Taxonomic information Pinto-Escobar 1976,1985; and Blanco & del Valle on Festuca in the paramo zone of South America 1991). Seven species are present in Humboldt's (Venezuela, Colombia, Ecuador, N Peru, SE Bra- collection, five of which were later described by zil) has been quite sporadic, although botanical C.S. Kunth as new (F. andicola, F caldasii, F. dasy- collections have increased in the last 50 years. antha, F procera, F. quadridentata). The Festuca The oldest specimens of Festuca from Venezuela specimens collected by T. Haenke (Expedicion were collected during 1906-1911 by A. Jahn, Malaspina 1789-1794) are of even earlier date, but H. Pittier. The first list of Venezuelan grasses was none of them are from Ecuador or Colombia. The published in 1937 (Anonymous 1937) and only uncertain origin of the type specimen of Festuca two species of Festuca (F. livida and F tolucensis) presliana Hitchc. (= Bromus depauperatus J. Presl) were listed. In 1942, Luces distinguished between is discussed in Stancik & Peterson (2003). the Mesoamerican and the Venezuelan popula- tions of F. livida and described a new Venezuelan Hitchcock (1927), in his overview of grasses species, Ffragilis (Luces) Briceno. Two species of Ecuador, Peru, and Bolivia, mentioned 17 spe- {Ffragilis and F tolucensis) were mentioned in the cies. The Ecuadorian species included^ dasyantha, ' Department of Botany, Charles University of Prague, Bendskd 2, 128 01 Prague 2, Czech Republic, [email protected] 2 Department of Botany MRC-166, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA. [email protected] Festuca in South American Paramos F. ulochaeta, F. eminens, F procera, F sublimes, F hatico, F monguensis, F nereidaensis, F pilar- F. andicola, F. dichoclada, F. dolichophylla, franceii,