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Memoirs of The New York Botanical Garden ADVISORY BOARD PATRICIA K. HoLMGREN, Director of the Herbarium The New York Botanical Garden JAMES L. LUTEYN, Senior Curator The New York Botanical Garden
ScoTT A. MoRI, Senior Curator The New York Botanical Garden EDITORIAL BOARD
WILLIAM R. BucK, Editor The New York Botanical Garden Bronx, New York 10458-5126 WM. WAYT THOMAS, Associate Editor The New York Botanical Garden Bronx, New York 10458-5126 THOMAS F. DANIEL (1991-2001) Department of Botany California Academy of Sciences Golden Gate Park San Francisco, California 94118
The MEMOIRS are published at irregular intervals in volumes of various sizes and are designed to include results of original botanical research by members of The New York Botanical Garden's staff, or by botanists who have collaborated in one or more of The New York Botanical Garden's research programs. Ordinarily only manuscripts of lOO or more typewritten pages will be considered for publication. Manuscripts should be submitted to the Editor. For further inforrnation regarding editorial policy and instructions for the preparation of manuscripts, contact the Editor. Orders for published and forthcoming issues and volumes should be placed with:
Scientific Publications Department THE NEW BOTANICAL GARDEN Bronx, New York 10458-5126 U.S.A. (718) 817-8721; fax (718) 817-8842 email: scipubs@nybg.org
a checklist ofplant diversity,
geographical distribution, andbotanical literature
James L. Luteyn with contributions from
Steven P. Churchill Dana Griffin III S. Rob Gradstein Harrie J. M. Sipman Mauricio R. Gavilanes A.
Memoirs ofThe NewYorkBotanical Garden Volume 84
© Copyright 1999 by The New York Botanical Garden Press
Published by The N ew York Botanic~l Garden Press
The páramo scene on the cover is taken from a mural that hangs in The Field Museum (Chicago). The mural was painted in 1948 by staff artist Arthur. G. Rueckert from photographs and other material provided by José Cuatrecasas. This mural shows a scene from "one of the most beautiful and least visited páramos ofColombia"-·the Sierra Nevada del Cocuy (compare with Fig. 4). The dominant species depicted in the mural is Espe/etia /opezii. (Reproduced with perrnission from The Field Museum, Chicago [Neg #B80317].)
The paper used in this publication meets the requirements ofthe American National Standard for Information Sciences-Perrnanence of Paper for Publications and Documents in Libraries and Archives, ANSI/NISO (Z39.48-1992). Printed in the United States of America using soy-based ink on recycled paper
Líbrary of Congress Catalogíng-in-Publicatíon Data Luteyn, James L. Páramos : a checklist of plant diversity, geographical distribution, and botanicalliterature 1 James L. Luteyn; with contributions from Steven P. Churchill ... [et al.]. p. cm.- (Memoirs ofthe New York Botanical Garden; v. 84) Includes bibliographical references. ISBN 0-89327-427-5 l. Paramo plants--Andes Regían. 2. Paramo ecology-Andes Region. 3. Paramos-Andes Region. I. Title. II. Series. QK1.N525 vol. 84 [QK241] 580 s----dc21 [581.7'53'098] 99-17171
"!
1
Contents Abstract Dedication Foreword • S.A. Mari Contributors Preface Color Plates Introduction to the Páramo Ecosystem General Definition Geographical Distribution Clima te Soils Paleohistory and Paleoecology Vegetation Zonation Superpáramo Grass Páramo Subpáramo Morphological and Physiological Adaptations Growth Forms Rosette Plants Cushion Plants Bunchgrasses Microphyllous Shrubs Prostrate Dwarf Shrubs Geophytes Vagrant Plants Flora General Floristic Diversity General Phytogeography and Origins Fauna Hurnan Influence The Pre-Columbian Period The Colonial and Independence Periods The Modem Period Impact of Buming and Grazing Vegetative Structure, Composition, and Species Diversity
Vlll
ix X XI Xll XVll
1 1 2 3
4 6 9 10 10 17 19 20 21 23 23 23 24 24 25 25 25 28 30 30 32 32 33 34 34
Soil Structure and Water Storage Capacity Decomposition, Nutrients, and Productivity The Future of the Páramo Ecosystem Scientific Reasons Ecological Reasons Economic Reasons Management Conservation Environmental Education
35 36 36 36 37 37 38 38 38
Checklist of Páramo Plants Lichens • H. J M Sipman Lichenized Ascomycetes Lichenized Basidiomycetes Mosses • S. P. Churchill & D. Griffin !JI Hepatics • S. R. Gradstein Anthocerotae Hepaticae Vascular Plants • J L. Luteyn Fems and Fem Allies Gymnosperms Angiosperms
41 41 42 51 53 65 65 66 74 76 82 82
137
Gazetteer of Páramo Localities Costa Rica and Panama • J L. Luteyn Colombia • J L. Luteyn & M R. Gavilanes A. Venezuela • J L. Luteyn & M R. Gavilanes A. Ecuador • J L. Luteyn & M. R. Gavilanes A. Peru • J L. Luteyn Bibliographic Sources for Locations ofPáramos
138 142 172 194 220 226
Botanical References to Páramo
229
Appendices Appendix 1: Families and Genera in the Páramo Checklist Appendix 2: Nomenclatura! Novelties
272 278
The New York Botanical Garden Press thanks the following donors for their generous support of this publication:
The Andrew W. Mellon Foundation Metropolitan Life Foundation, a leadership funder of The New York Botanical Garden's Scientific Publications Program.
Pรกra111os
ABSTRACT Luteyn, J. L. (Institute of Systematic Botany, The N ew York Botanical Garden, Bronx, NY 10458-5126, U.S.A.). Páramos: A checklist ofplant diversity, geographical distribution, and botanicalliterature. Memoirs ofThe New York Botanical Garden 84: 1-000. 1998.-Páramo is restricted to the northem Andes ofSouth America and adjacent southem Central America. It is a high-elevation ecosystem, from ca. 3000 m to 5000 m, between the continuous forest line and the upper limit of plant life at the snowline. It is characterized by tussock grasses, large rosette plants, shrubs with evergreen, coriaceous, and sclerophyllous leaves, and cushion plants. It has high biological diversity and is rich in endemism, but it is ecologically fragile. Páramo is a center of economic importance in the high Andes, .especial! y for agriculture and cattle raising. Its most important role, however, relates to Andean hydrology. Páramo has the richest high-mountain flora in the world. This book includes the family, genus, authority,·pertinent synonymy, and geographical and altitudinal ranges for each ofthe 3399 vascular and 1298 nonvascular páramo plant species. It a,lso gives the country, proper name, maximum altitude, latitude and longitude, political subdivisions, and notes for approximately 2100 páramo localities, plus fullliterature citations for more than 1570 páramo botanical references. Many ofthe black-and-white photos illustrating the book were taken by one ofthe early students ofpáramo, José Cuatrecasas.
'Key words: Páramo, South America, Central America, Andes, mountains, ecology, vegetation biogeography, human influence, taxonomy, checklist, lichens, mosses, hepatics, vascular plants, gazetteer, botanicalliterature.
RESUMEN Luteyn, J. L. (lnstitute of Systematic Botan y, The New York Botanical Garden, Bronx, NY 10458-5126, U.S.A.). Páramos: A checklist ofplant diversity, geographical distribution, and botanicalliterature. Memoirs ofThe New York Botanical Garden 84: 1-000. 1998.-El páramo está restringido al norte de los Andes de Sur América y a las áreas adjacentes al sur de Centro América. Es un ecosistéma de alta montaña, el cual va desde 3000 m a 5000 m de altitud, entre la franja de bosque continuo y el límite superior de nieve perpetua donde aún ocurre vida vegetal. Está caracterizado por gramíneas en macolla, plantas en rosetas, arbustos siempreverdes con hojas coriáceas y esclerófilas, y plantas en cojín. Tiene una diversidad biológica alta y es rico en endemismo, pero es ecológicamente frágil. El páramo es un centro de importancia económica en los altos Andes, especialmente en agricultura y ganadería. Sin embargo, su papel más importante está relacionado con la hidrología andina. El páramo tiene la flora más rica de alta montaña en el mundo. Este libro incluye familia, género, especie, autor, sinónimos pertinentes y rangos geográficos y altitudinales para cada una de los 3399 especies de plantas vasculares y 1298 especies de plantes no vasculares del páramo. También presenta el país, el nombre proprio, la altitud máxima, la latitud y la longitud, las divisiones políticas y algunas notas sobre aproximademente 2100 localidades de páramo, más citaciones de literatura completa para más de 1570 referencias botánicas del páramo. Muchas de las fotos en blanco y negro que ilustran el libro fueron tomadas por uno de los estudientes pioneros del páramo, José Cuatrecasas. Palabras clave: Páramo, América del Sur, América Central, Andes, montañas, ecología, vegetación, biogeografía, influencia humana, taxonomía, lista anotada, líquenes, musgos, hepáticas, plantas vasculares, diccionario geográfico, literatura botánica. viii
DEDICATION I am proud to dedicate this book to the memory of two outstanding field botanists, scholars, gentlemen, and friends-José Cuatrecasas and Luis Ruiz-Terán. José Cuatrecasas Arumí (1903-1996) (Fig. 1), "Don José," is tome the father ofmodern páramo studies. Born in Spain, he made his first trip to Colombia in 1932. His lo ve for the neotropical flora in general and for the páramos of Colombia in particular is reflected in the large number of botanical collections he made, the numerous taxonomic revisions and general papers he published (see Robinson et al., 1996), and the excellent photographs he took. His charm (e.g., while entertaining at home, showing slides and eating Chinese takeout), friendship, and many years of dedicated service to botany willlong be remembered by those of us who knewhim. Luis Enrique Ruiz-Terán (1923-1979) (Fig. 2) was born and raised in the páramo region ofMérida, Venezuela. He maintained a humble lifestyle, dedicating mostofhis energy to the study ofbotany. His special !ove for páramo is reflected in the great number ofvery detailed and carefully documented herbarium collections he made. As a professor ofbotany at the Universidad de Los Andes in Mérida, he influenced and stimulated many ofus to appreciate the páramo. Those ofus who knew him in the field (where he frequently stopped to prepare "tinto" and smoke a cigarette) will never forget the man. He did not publish much, but his intimate knowledge of and dedication to the flora of his native Andes was unsurpassed. Luis was also an avid mountain climber, and Pico Ruiz-Terán was named after him.
Fig. l. José Cuatrecasas standing alongside a population of Coespeletia timotensis at 4200 m in Páramo de Piedras Blancas (Mérida, Venezuela). (From Rundel et al., Tropical Alpine Environments, Cambridge University Press, New York. Reproduced with permission from Cambridge University Press. Photo probably taken by AJan P. Smith in about 1978.)
Fig. 2. Luis Ruiz-Terán standing next to his namesake, Ruilopezia ruizii, at ca. 2800 m in Páramo de Las Coloradas (Mérida, Venezuela). (Photo Cuatrec. I-4420, probably taken by José Cuatrecasas in about 1978.) ix
FOREWORD The Andes harbor the richest assemblage ofplants and animals in the Neotropics and perhaps in the world. The wide range ofhabitats caused by vast differences in elevation (from sea level to nearly 7000 meters at the summit of Aconcagua in Argentina), enormous differences in rainfall (from no rain in a given year in the Atacama Desert to 4500 mm per year at the foothills ofthe Andes in westem Amazonia), and the relative isolation of numerous mountain valleys have combined to produce ni ches inhabited by countless plant and animal species. The crown jewel of Andean vegetation types is páramo, located abo ve the last continuous forests and below the permanent snowline between 8°S and 11 °N latitudes in Venezuela, Colombia, and Ecuador, but with a few outliers in northem Peru in the south and' Panama and Costa Rica in the north. Páramo plays and has played an essential role in the ecology and evolution of Andean ecosystems. Despite its relatively small surface area-no more than 2% of the land area of the countries in which it is foundthe páramo flora is nevertheless extremely diverse with many endemic species. In fact, it is the richest high-mountain flora ofthe world (Smith & Cleef, 1988). Within the confines ofpáramo, Jim Luteyn and his collaborators have registered 3399 vascular and 1298 nonvascular plant species. Today, as in the past, páramo also acts as a center of economic importance in the high Andes, especially for agriculture and cattle raising. Its most importantrole, however, relates to Andean hydrology, where it acts as a sponge absorbing moisture from rain and fog for subsequent use by plants and animals, including man, in all of the Andean ecosystems. The expansion and contraction of páramo in conformity with climatic changes in the Pleistocene had a tremendous impact on the evolution and subsequent distribution of plants and animals. During glacial periods páramo expanded into lower elevations, and during interglacial periods it contracted to higher elevations. These movements created the opportunity for evolution ofboth páramo and lower-elevation plant and animal species. Jim became interested in páramo in 1978, on his first trip to Venezuela, where he was fascinated by its general beauty, interesting life forms, and variety of plants, including those he studies, the N ew W orld blueberries (Ericaceae ). The blueberries are the seventh largest family of flowering plants in the páramo (see Table VI), currently known to be represented by 16 genera and 79 species. During that trip Jim met José Cuatrecasas and Luis Ruiz-Terán for the frrst time, and formulated the idea (with Ruiz-Terán) of a revision ofVareschi' s (1970) Flora de Los Páramos. Since then Jim has retumed to the Andes nearly 50 times, and on many of those trips he collected in páramo. In 1992, Jim Luteyn and Henrik Balslev edited a multi-authored book entitled Páramo: An Andean Ecosystem under Human Injluence, following a symposium they organized in Aarhus, Denmark. This current book is, therefore, the result oflong-term interests Jim and his collaborators have had in páramo. This carefully crafted reference book to the páramo ecosystem will serve as a basis for a future páramo flora. It is divided into four parts: an introduction to páramo, checklists of plants, a gazetteer of páramo localities, and references to páramo in the botanicalliterature. Each part provides basic information for those interested in practically any aspect ofpáramo. For example, professors teaching courses in plant ecology will want to consult the introduction for the detailed descriptions of the different plant growth forms found in páramo; botanists producing local floras will need to refer to the checklists of lichens, mosses, hepatics, and vascular plants; geographers will fmd a great deal of information in the gazetteer enabling them to locate places where páramo plants have been collected, thereby facilitating the production of maps showing geographic distributions; the detailed botanical reference section will be indispensable to anyone interested in Andean vegetation; and conservationists and park managers will find data useful to them in all parts of the book. The strength of this book is that, for the first time, it assembles in one place information covering the entire páramo ecosystem that ranges from Costa Rica to Peru, and includes all higher plant groups, inX
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cluding lichens, mosses, hepatics, and vascular plants. Moreover, the data has been derived from an exhaustive search of numerous Latin American, North American, and European libraries and herbaria. The checklists provided by Churchill, Griffin, Gradstein, and Sipman have added data for groups of their expertise. For botanists interested in Andean plants this book is an essential reference. The checklist, in lieu of a complete flora, will facilitate plant identification. Although most botanists have a general idea ofthe classification of plants they observe and collect, they frequently need a memory aid in the form of a list of names to allow them to go to the right places in herbaria to confirm their determinations. Moreover, future botanists wishing to produce guides to the lichens, mosses, hepatics, and vascular plants of páramo have in this book the first step, a checklist, already done for them. Many ofthe photos illustrating the book were taken in the 1930s and 1940s by one ofthe early students of páramo, Dr. José Cuatrecasas, and depict páramo before much of it was disturbed by man. Today, similar! y undisturbed páramo is difficult, if not impossible, to find because of excessive human interference. Jim Luteyn and his collaborators have assembled the information upon which sound conservation plans can be developed to protect remaining páramos throughout the Andes. The authors will have achieved a very important goal ifthis book stimulates govemments and conservation organizations to preserve and restore representative páramos to the state they were observed by José Cuatrecasas and Luis Ruiz-Terán, pioneer páramo botanists to whom this book is dedicated.
Scott Mari The New York Botanical Garden, 1999
CONTRIBUTORS Steven P. Churchill- Institute of Systematic Botany, The New York Botanical Garden, Bronx, New York 10458-5126, U.S.A.; current address: Dept. 23, Missouri Botanical Garden, Box 299, St. Louis, Missouri 63166-0299, U.S.A. Mauricio R. Gavilanes A.- Av. Amazonas 1230 y Foch, Local "ARAMIS," Quito, Ecuador. S. Rob Gradstein - Systematisch-Geobotanisches Institut, University of Gottingen, Untere Karspüle 2, 37073 Gottingen, Germany. Da na Griffin IH- Florida Museum ofNatural History, University ofFlorida, Gainesville, Florida 32611-2035, U.S.A. James L. Luteyn- Institute ofSystematic Botany, The New York Botanical Garden, Bronx, New York 10458-5126, U. S.A. Harrie J. M. Sipman- Botanischer Garten und Botanisches Museum Berlin-Dahlem, KoniginLuise-Stra!3e 6-8, D-14191 Berlin, Germany.
PREFACE Above the montane forest and below the permanent snowline, a mostly open vegetation collectively known as páramo is scattered within the humid equatorial Andes. Like isolated mountaintops, islands in the sky, the páramo ecosystem is discontinuously distributed between 11 °N and 8°S latitudes, primarily in Venezuela, Colombia, and Ecuador, but also with a few outliers to the north in Costa Rica and Panama and to the south in northern Peru. The páramo ecosystem has high biological diversity and is rich in endemism, but it is ecologically fragile. Though it has provento be resilient to human disturban ce, it is rapidly reaching its limits. With human population size at its greatest in history and steadily rising, pressure for increased food production and land use within páramo has intensified, and deterioration of the environmenf has inevitably followed. With the public' s increased desire for leisure time, there is also a new awakening to the potential of the páramo for recreation and tourism. Therefore, the páramo ecosystem is in a precarious environmental position and is being threatened. There is an urgent need to in crease our efforts to study this ecosystem, not only to know what is there by means of inventory and ecological analyses, but also to monitor the effects of disturbance upon the vegetation and aquatic resources. Páramos have fascinated scientists ever since the first Europeans visited them in South America. Plants from páramo were first collected by Mutis in Colombia in the late 1700s and formally described by Humboldt (1816, 1817 for Colombia), Goebel (1891 for Venezuela), and Diels (1937 for Ecuador). In the late 1800s, Funck and Schlim, Jameson, Karsten, Purdie, H. H. Smith, Spruce, Triana, and others collected páramo plants. In the first half of the 20th century, Killip and A. C. Smith, Troll, Cuatrecasas, and Jahn were actively collecting, and more recently Acosta-Solís, H. Barclay, Ewan, Fassett, Fosberg, Haught, Little, López-Figueiras, Ruiz-Terán, Wm. Steere, Steyermark, and Vareschi, among others, gathered large numbers of páramo plants. In recent years, the Universidad de Los Andes (Mérida, Venezuela) has become a leading center ofpáramo ecology through its Centro de Investigaciones Ecológicas de los Andes Tropicales (CIELAT). Also, the Instituto de Ciencias Naturales (Universidad Nacional de Colombia, Bogotá) and the Instituto Geográfico 'Agustín Codazzi' (Bogotá), in cooperation with the Hugo de Vries Laboratory (University of Amsterdam, The Netherlands), has significantly advanced our knowledge about the paleoecology and community structure of páramo in Colombia through its ECO ANDES program (Estudios de Ecosistemas Tropandinos). It is because of all this activity, associated with many collecting efforts, that there are literally thousands ofherbarium specimens of páramo plants available for study. Therefore, from a taxonomic and ecological point of view, certain aspects of páramo have been relatively well studied. Currently, however, there is no written flora ofthe entire páramo, and the only means to identify páramo plants is by using individual, widely scattered monographs or comparisons in the herbarium. Neither is there any authoritative book that summarizes our overall knowledge about páramo or guides one to literature about this ecosystem. Furthermore, although the vascular páramo flora is one ofthe richest in genera (500) and species (3399) ofthe high mountains of the world, the cold montane tropics are still overlooked by students of tropical biology, who focus more on the warm lowlands (Vuilleumier & Monasterio, 1986). My own interest in páramos started in 1978-1979, with m y first collecting trips to Venezuela, in the company ofLuis Ruiz-Terán. After working together for nearly two years, we decided to begin a revision ofVareschi's (1970) Flora de los Páramos. Unfortunately, this idea was cut short by Luis's unexpected death in 1979. My own interest remained dormant, although I have always tried to collect in the páramos on each field trip to northwestern South America. In 1986, on a trip to the Netherlands, I met with Antaine Cleef and we put together an idea for a generic flora and guide to the páramos in collaboration with Orlando Rangel Ch. (from Colombia). We had four purposes for xii
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the project: 1) to encourage and in crease botanical research in the páramo ecosystem; 2) to publish a checklist ofthe moss, hepatic, lichen, and vascular plant species; 3) to publish in Spanish an illustrated manual ofthe genera ofvascular plants suitable for both the layman and specia1ist; 4) to increase pub1ic awareness of páramo with the aim of stimulating efforts to conserve the páramo ecosystem. The fact that páramo areas have received re1atively good attention from botanists (compared to lowland and even sorne middle c1oud-forest habitats) made a páramo flora project feasib1e. The idea ofthe páramo project was formally presented at the first Páramo Symposium in Aarhus, Denmark in 1991 (Luteyn et al., 1992). Later I decided to put together a database of information necessary to begin such a flora, enlisting the help of specialists in lichens, hepatics, and mosses. In 1994 1 began to construct three databases: one that included for each páramo plant species its family, genus, authority, synonymy, geographical and altitudinal ranges, páramo subdivision, voucher specimen, and notes; a second database for each páramo locality giving its country, proper name (sometimes cross-referenced to other names used), maximum altitude, latitude and longitude, politica1 subdivisions, and notes; and a third database for botanical1iterature. These three databases form the basis for the current volume, which is the first major result ofthat idea born in 1979. In 1996, for practica! reasons, I rea1ized that I would not be able to complete a flora; therefore, I have brought together the current book to "pave the way" and stimu1ate the production of a full flora ofthe páramos. The emphasis ofthis book is taxonomic; details of ecological, evolutionary, and biogeographical studies should be sought elsewhere. There is very litt1e 1iterature about the páramos of northern Peru, and relatively little about those of Ecuador and Costa Rica; therefore, this vo1ume' s "Introduction to the Páramo Ecosystem" is based mostly on the páramos of Venezuela and Colombia, where the 1iterature abounds. Because there has not been any general flora of the páramos, nor for any high montane region in Latin America since Weddell (1855), this book will help fill a 1arge gap in our lmowledge about the páramo ecosystem and hopefully will become a standard reference for the high-e1evation flora ofthis region. Not only should it stimulate further work in the páramo-enabling comparisons ofthe richness, distribution, evolution, and biogeography of different taxa-but it should a1so provide the necessary raw data often requested by scientists, pub1ic officials, and_nongovernmental agencies to intelligently address conservation issues. While the check1ist provides important floristic information, it is nota substitute for a written flora ofthe páramos, which is still sorely needed. Finally, as with so many other experiences in life, it may be difficult for sorne to understand and appreciate the páramo solely from this book, if it has not already been experienced firsthand. Therefore, the book begins with a brief description ofpáramo: its definition, general ecology, life forms, growth habits, vegetation, and conservation. Once páramo has been experienced personally, this book will become much more meaningful and the páramo easier to visualize.
Acknowledgments.-There are so many people to whom I need to say "thank you" for helping me with the many different aspects ofthis book. Ifi have forgotten anyone, it is certainly unintentional and I do appreciate your efforts. 1 showed sorne stage of the manuscript to many peop1e, especially specialists in different plant groups, whom I have listed alphabetically by the plant family in which they assisted: Acanthaceae (D. Wasshausen), Alstroemeriaceae (R. Gereau, M. Neuendorf), Apiaceae (L. Constance), Asclepiadaceae (G. Morillo), Asteraceae (A. Anderberg, V. Badillo, J. Cuatrecasas, S. Díaz P., M. Dillon, V. Funk, B. Nordenstam, J. Panero, J. Pruski, H. Robinson), Berberidaceae (C. Ulloa U.), Brassicaceae (I. Al-Shehbaz), Bromeliaceae (J. Betancur), Buddlejaceae (E. Norman), Campanulaceae (T. Ayers, T. Lammers), Chloranthaceae (C. Todzia), Clethraceae (C. Gustafsson), C1usiaceae (N. Robson), Crassulaceae (A. Freire-Fierro), Cyperaceae (A. Reznicek, M. Strong, W. Thomas, G. Tucker, G. A. Wheeler), Eriocaulaceae (N. Henshold), Fabaceae (R. Barneby, J. Grimes), ferns
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(D. Lellinger, B. León, J. Mickel, R. Moran, A. R. Smith), Gentianaceae (J. Pringle), Gesneriaceae (L. Skog), Juncaceae (H. Balslev), Lamiaceae (R. Hariey), Liliaceae (R. Cruden), Loasaceae (M. Weigend), Loranthaceae s.l. (J. Kuijt), Lycopodiaceae (B. 0llgaard), Malvaceae (L. Dorr, P. Fryxell), Melastomataceae (F. Almeda, S. Renner, J. Wurdack), Myrsinaceae (J. Pipoly), Myrtaceae (L. Landrum), Onagraceae (P. Berry, J. Solomon), Orchidaceae (K. Barringer, E. Christenson, C. Dodson, G. Romero), Piperaceae (R. Callejas), Poaceae (B. Briceño, L. Clark, G. Davidse, S. Lcegaard, P. Peterson, S. Renvoir), Polygalaceae (B. Eriksen, J. Wurdack), Potamogetonaceae (R. Haynes), Rosaceae (K. Romoleroux), Rubiaceae (J. Kirkbride), Scrophulariaceae (F. Astholm, J. L. Fernández-Alonso, N. Holmgren, U. Molau), Selaginellaceae (1. Valdespino), Solanaceae (C. Benitas de Rojas, S. Knapp, M. Nee, C. Ochoa), Symplocaceae (B. Stahl), Valerianaceae (B. Eriksen), Violaceae (U. Molau). I also want to express my gratitude to the library and/or herbarium staff ofthe following institutions for allowing access to their collections andhelp iri finding materials: The New York Botanical Garden (Bronx), Smithsonian Institution (Washington, DC), Institute of Systematic Botan y at the State UniversitYofUtrecht and the Hugo de Vries L<tboratory ofthe University of Amsterdam (the Netherlands), Department ofSystematic Botany ofthe University of Aarhus (Denmark), Botanical Museum (Goteborg) and Herbarium of the Swedish Museum of Natural History (Stockholm), Instituto de Ciencias Naturales ofthe Universidad Nacional de Colombia (Bogotá), Universidad de Los Andes (Mérida, Venezuela), Herbario. Nacional de Venezuela (Caracas), Herbario de la Facultad de Agronomía de la Universidad Central de Venezuela (Maracay), and the Instituto Venezolano de Investigaciones Científicas, IVIC (Caracas), Herbario QCA ofthe Pontificia Universidad Católica del Ecuador and Museo Nacional de Ciencias Naturales (Quito), Missouri Botanical Garden (St. Louis), Field Museum (Chicago), Harvard University Herbaria (Cambridge), Royal Botanic Gardens (Kew), Herbario Nacional de Costa Rica (San José), and the American Museum ofNatural History and New York Public Library (New York). I want to extend m y appreciation to the Department ofBotany, Smithsonian Institlition, for awarding mean Andrew W. Mellon Foundation "Senior Mellon Fellowship" (under the sponsorship of Laurence J. Dorr) during 1993-1994, to work at the United States National Herbarium (Washington, DC) to begin compiling the data. It was during m y tenure there that I was really able to get this project off the ground. They are also to be thanked for allowing me free access to the late José Cuatrecasas's office, where I was able to search for and select from don José's original negatives of páramo habitats and plants that so embellish the pages ofthis book, and to see his final unpublished manuscript "A Systematic Study ofthe Subtribe Espeletiinae (Heliantheae, Asteraceae)." Thanks also to the Smithsonian Institution Center for Scientific Imaging and Photography for their care in printing the negatives, sorne of which were glass negatives from the 1930s. Cuatrecasas 's photos are cited by his abbreviated name followed by the series and negative number (e.g., "Cuatrec. C-276"). M y thanks al so go to the following individuals who helped me in sorne additional way: J. Aguirre, G. Aymard, C. Benitas de Rojas, P~ Berry, J. Betancur, F. Bouman, B. Briceño, B. Buck, P. Buck, R. Callejas, E. Cotton, S. Churchill, T. Croat, D. Díaz-Miranda, L. Dorr, E. Farr, J. L. FernándezAlonso, J. Fernández Casas, L. D. Gómez P., F. González, R. Gradstein, D. Griffin III, Th. Van der Hammen, R. Hofstede, L. Hollenberg, H. Hooghiemstra, S. Horn, O. Huber, M. Kappelle, P. Keating, S. León, Y. León, J. F. Morales, S. A. Morí, G. Morillo, J. Mulroy, F. Pérez, T. Pocs, G. Raeymaekers, B. Ramírez P., P. Ramsay, O. Rangel Ch., R. Riina, H. Rodríguez, A. Sagástegui A., S. Salamanca V., D. Sánchez, l. Sánchez Vega, H. Sipman, P. Sklenáf, J. Solomon, H. Witte, and K. Young. 1 thank Bobbi Angell for preparing the maps and climatic diagrams. Special thanks go to those who were especially generous with their time and efforts: H. Balslev andA. M. Cleef for conversation, reading and correcting manuscript, and support over the years; M. Gavilanes A. for hclp in all aspects ofthe initial stages ofthe checklist and gazetteer; F. Sarmiento and C. Ulloa U. (Ecuador) and G. Morillo and B. Briceño (Venezuela) for careful reading and helpful comments, especially on the
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gazetteers; J. Sánchez G. for allowing me access to his (and G. Vargas Ulate) unpublished manuscript on the páramo plants of Costa Rica and for helping with the Costa Rican gazetteer; P. M. J0rgensen for helpful comments and sharing personal data from his experiences in Ecuador, and along with the MissouriBotanical Garden for sharing data from TROPICOS; l. Sánchez Vega for sharing his checklist ofplants from the Peruvianjalca; M. Sánchez Montoya for gathering information for the gazetteer ofPeru; and to F. González, O. Huber, S. Mori, C. Peters, and K. Young for taking so much oftheir time toread the manuscript in its final stages and for their helpful comments and constructive criticisms; andan extra special thanks to D. S. Sylva S. for helping with computerization, reading and correcting text, and encouraging me in so many other ways throughout this project. In addition, 1 want to thank all those colleagues, students, guides, and "mulleteros" who accompanied me on various trips into the páramo, and to the many campesinos who gave considerable assistance by means of their kindness and access to their homes and food; the only things warmer than their hospitality were the many bottles of rum and aguardiente that we shared during the very cold nights! One last word: This book is the result of a labor oflove for the páramo ecosystem, through which 1 have so often passed and in which 1 have spent so many enjoyable days collecting plants. 1 have tried to pull together in this one volume as much information as possible about all botanical aspects of páramo, in the hopes that this effort will stimulate and encourage others to study the páramo and to publish a full flora of this marvelous ecosystem.
James L. Luteyn New York, 1999
COLOR PLATE I. Top left, Páramo Las Cajas (Cuenca, Ecuador), 4300 m. Note thejagged terrain with numerous glaciallakes and grass páramo. (Photo: P. Keating, taken 1991.) Top right, Jalea de Las Lagunas (Cajamarca, Peru), 4500 m. Glaciallakes surrounded by Stipa-dominated, short-grass páramo. (Photo: J. L. Luteyn, taken 1995.) Bottom left, Páramo de Mojanda (Imbabura, Ecuador), 3 700 m. Grass páramo dominated by Calamagrostis spp. with Polylepis sp. forest growing along steep slopes and cliffs. Note the sharp boundary between the forest and páramo, probably dueto the influence offire. (Reproduced with permission ofthe Department ofSystematic Botany, University of Aarhus, Denmark. Photo: B. 011gaard, taken 1976.) Bottom right, Volcán Cotopaxi (Cotopaxi, Ecuador), 4000 m. Bunchgrass páramo dominated by species of Festuca and Calamagrostis. (Photo: J. L. Luteyn, taken 1997.)
CoLOR PLATE II. Top left, Páramos de Matanga (Azuay, Ecuador), 3300 m. Mosaic of grass páramo and isolated patches offorest. (Photo: C. Ulloa U., taken 1990.) Top right, Volcán Chiles (Carchi, Ecuador), 4200 m. Páramo bog with mats of Distichia muscoides and Plantago rigida. (Photo: J. L. Luteyn, taken 1989.) Bottom left & right, Páramos del Angel (Carchi, Ecuador), 3500 m. Grass páramo dominated by the bunchgrass Calamagrostis effusa, the stem rosette Espeletia pycnophylla, and the acaulescent rosette Puya hamata. (Photos: bottom left, H. Ba1slev, taken 1995; bottom right, P. M. Jorgensen, taken 1983.)
CoLOR PLATE Ill. Top left, Páramo de Piñango (Mérida, Venezuela), 3500 m. Each October this páramo is decorated with magenta flowers of Chaetolepis lindeniana and yellow flowers ofEspe/etia schultzii and Hypericum /aricifolium. (Photo: J. L. Luteyn, taken 1996.) Top right, Páramo de Mucubají (Mérida, Venezuela), 3500 m. Steep slopes showing yellow inflorescences ofEspeletia schultzii and fields ofimmaturewheat with red patches ofRumeX: acetosel/a, an aggressive colonizerofold fields. (Photo: J. L. Luteyn, taken 1996.) Bottom left, Volcán Antisana (Pichincha, Ecuador), 4000 m. The gradual transition frumforest belowtopáramo above. Dominants are Polylepis incana, Gynoxys acostne, Baccharis sp., Loricaria thuyoides, and L. antisanensis. (Photo: J. L. Luteyn, taken 1992.) Bottom right, Volcán Chimborazo (Chimborazo, Ecuador), 4200 m. Desert páramo with drifting and eroding sand. Plants commonly scattered include the bunchgrass Stipa, low shrubs of Chuqiraga, and circular clumps ofBidens (yellow flowers shown). (Photo: J. L. Luteyn, taken 1997.)
CoLOR PLATE IV. Top left, Senecioformosus, a common plant in the Páramo de Piedras Blancas above lv!érida (Venezuela), which provides spectacular displays ofmagenta flowers in October and November. (Photo: J. L. Luteyn, taken 1996.) Top right, Gentianella hyssopifolia, a colorful herb from the páramos south ofCuenca (Ecuador). (Photo: J. L. Luteyn, taken 1990.) Bottom left, Ca/ceo/aria ericoides, a suffrutescent herb found in the shelter of rocks on Volcán Chimborazo (Ecuador). (Photo: J. L. Luteyn, taken 1997). Bottom right, Jamesonia cinnamomea, a "cobra" fem in the grass páramo on Volcán Cayarnbe (Ecuador). The dense reddish-brown pubescence and curled leaves help protect the plants from the harsh environment. (Reproduced with permission ofthe Department of Systematic Botany, University of Aarhus, Denmark. Photo: L. B. Holm-Nielsen.)
CoLOR PLATE V. Top left, Loricaria thuyoides, from upper grass páramo on Volcán Chiles (Ecuador), is a microphyllous shrub with small, sclerophyllous, imbricate leaves on flattened stems. (Photo: J. L. Luteyn, taken 1989.) Top right, Festuca dolichophylla, a characteristic bunchgra~s in this grazed páramo on Volcán Cotopaxi (Ecuador). Note the bigh proportion of dead shoots among the living, which gives grasslands their silvery-grayish or brownish look and provides insulation for buds and young leaves. (Photo: J. L. Luteyn, taken 1997.) Bottom left, Bleclmum schomburgkii, a population oftree fems in the mist on Volcán Sumaca (Ecuador) showing the stem rosette growth form. (Photo: Department ofBotany, University of Aarhus.) Bottom right, Valeriana rígida, a small plantwith white flowers from Páramo Las Cajas (Ecuador) that combines sharply pointed, rigid leaves with an acaulescent rosette/cushion growth form. (Photo: J. L. Luteyn, taken 1990.)
CoLOR PLATE VI. Top left, Xenophyllum humile, a hemispherical, cushion plant from the Páramo de Guamaní (Ecuador). (Photo: J. L. Luteyn, taken 1981.) Top right, Orthrosanthus chimboracensis, an iris-like herb with succulent rhizomes from the Llanos de Cuibá (Colombia). An example of a plant with the geophyte growth form. (Photo: J. L. Luteyn, taken 1984.) Bottom left, Lupinus sp., a common herbaceous legume growing in the grass páramo ofthe Páramos de Matanga (Ecuador). (Photo: J. L. Luteyn, taken 1990.) Bottom right, Semiramisia pulcherrima, a scandent, climbing epiphytic shrub endemic to this Páramo de Bordoncillo region of southern Colombia. The orangeish-red, waxy flowers are up to 6 cm long and 2 cm in · diameter. (Photo: J. L. Luteyn, taken 1997.)
INTRODUCTION TO THE PARAMO ECOSYSTEM "The páramo zone is the most interesting of the life zones of the Andes, since it shows to the highest degree, the struggle ofplant and animallife against conditions of extreme cold temperature." [Chardón, 1933]
General Definition Within the tropical regions ofMexico, Central and South America, Africa, Malesia including New Guinea, and Hawaii, there is a vegetation type that occurs between the upper limit of continuous, closedcanopy forest (i.e., forest line or timberline) and the upper limit of plant Ji fe (i.e., snowline) that is characterized by tussock grasses, large rosette plants, shrubs with evergreen, coriaceous, and sclerophyllous leaves, and cushion plants. This vegetation type is §cattered along the crests of the highest mountain ranges or on isolated mountaintops from about 3000 m to 5000 m, like islands in a sea of forest. Locally these areas are lmown as "zacatonales" (the Mexican and Guatemalan volcanic highlands), "páramo" (Central and northem South America), "jalea" (northern Peru), "puna" (drier areas ofthe altiplano ofthe central Andes), "afroalpine" and "moorland" (East Africa), and "tropical-alpine" (Malesia). Although neither Beard (1944), Troll (1958b ), nor Lauer (1981) felt the term "alpine" was appropriate for intertropical high-altitude landscapes, these vegetation types have been more recently referred to as "tropical alpine" (Ramsay & Oxley, 1997; Rundel et al., 1994b; Smith & Young, 1987) or "tropicalpine" (Smith & Cleef, 1988). Ido not like to apply the general term "alpine" to vegetation in the tropics, because it is a term derived from temperate regions. Furthermore, high-elevation tropical climates differ sharply from those of tempera te alpine regions, particularly with respect to seasonality and diurna! pattems oftemperature change (Rundel, 1994). Sorne authors, including Walter (1973), Lauer ( 1981 ), and Monasterio and Vuilleumier (1986), advocate using the term "páramo" in the broadest sense possible, on a worldwide basis, for all high tropical montane vegetation abo ve the continuous timberline-for the sake of nomenclatura! simplicity, if for no other reason. Monasterio and Vuilleumierwould simply add a geographic adjective to characterize the particular area being discussed-Andean páramo, African páramo, and so forth. Andean páramo is often compared with
other alpine and arctic ecosystems (Baruch, 1979; Billings, 1973,1974, 1979;Billings&Mooney, 1968; Smith & Young, 1987). For the purposes ofthis book, the term "páramo" is used in its regional sense, being restricted to the northern Andes of South America and adjacent southem Central America. It is here used as a collective term for the entire landscape unit (or ecosystem) ofthe high altitude above continuous forest !ine and below the permanent snowline. Many different plant communities can be found in páramo, and will be discussed below, but the most widespread are dominated by tussock-forming grasses (Fig. 3). The word páramo comes from the Latín word "paramus," according to the Diccionario de la Lengua Española (Real Academia Española, 1992). Corominas (1973) states that this Hispanic-Latín word seemingly arose from the midwestern portion ofthe northem !berían Península but that its exact origin is uncertain. He further suggests that it may have been adopted by the Romans as a Celticism, or instead is neither Iberian nor Celtic but originates from another Indo-European language in pre-Roman times. In Spain, up until the epoch of the Conquest, the desertic platean of arid Castile, which contrasts with the fertile regions lower down, was called "paramera." Font Quer (1977) states that in Castilian Spanish the general significance ofthe word "páramo" is a flat plain. The early Spanish explorers applied the word "páramo" to north Andean areas that were high, cold, inhospitable, and wind- and rain-swept, perhaps reminding them of the plains of their native Iberian Península. In Colombia, atmospheric moisture in the form of drizzle is often referred to as "paramitos," while in Ecuador the term "parameando" has come to mean "it is raining," and in Venezuela one might say "estoy emparamado" when one is getting wet because of rain and cold (Monasterio, 1980b; Nuñes & Pérez, 1994; Ramsay, 1992; Vareschi, 1970; Weber, 1958; pers. comm. with local inhabitants). For definitions and insight into conceptualizations ofthe páramo ecosystem by local "parameros" (inhabitants ofthe páramos), see LópezZent, 1995, and Zambrano, 1993.
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Fig. 3. Páramo del Escoba! (Boyacá, Colombia) at ca. 3700 m elevation, dominated by the rosette plant Espeletia lopezii and grass Calamagrostis sp. with Hypericum mexicanum, Arcytophyllum sp. and Orthrosanthus chimboracensis. (Photo Cuatrec. C-1210A taken in 1938.)
The páramo landscape has been influenced by glaciation (Fig. 4). It is irregular and uneven, from jagged and very rough with erratic rocks to rolling or flat, oftentimes with many small glaciallakes and tributaries (Color Plates I-II). It is the source ofmany ofthe large rivers of northem South America (e.g., Río Magdalena and Río Cauca ofColombia, Río Napo and Río Coca of Ecuador, and Río Orinoco ofColombia and Venezuela). As will be seen from the discussions in this Introduction, there is no single definition of páramo, because it is characterized by a variety of geographic, geologic, climatic, physiognomic, and floristic features, all of which will be briefly touched upon.
Geographical Distribution In the tropics of the Americas, the páramo ecosystem is discontinuously distributed between 11 °N and sos latitudes. It is concentrated in the northwest corner of South America, mostly in Venezuela, Colom-
bia, and Ecuador, with sorne outliers in Costa Rica, Panama, and northem Peru (on the endpapers of this volume, the area above 3000 m elevation that is potentially páramo is shown in black). The northemmost páramos are located in the Sierra Nevada de Santa Marta of Colombia, located at about 11 °N. The westemmost páramos are located in Costa Rica, in the Cerro Buena Vista (Cerro de La Muerte) region of the Cordillera de Talamanca, at about 83°W longitude. The eastemmost páramos are located in north-central Venezuela, in the state of Lara, at about 70°W. The southernmost páramos (locally called "jalea") are found in northern Peru in the department of La Libertad, at about 8°S, just north of the Cordillera Blanca. Other neotropical areas that have páramo-like vegetation or that correspond ecologically and have sometimes been referred to as páramo, but that fall outside the geographical range or definition used in this book, include Pico Duarte in the Dominican Republic, the "zacatonales" of Mexico, Pico Naiguatá
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3
Fig. 4. Panoramic view ofNevado del Cocuy (Boyacá, Colombia) looking East from the Pozo Azul ranch at ca. 4110 m. The rocky peak to the left is Los Guasquines, the glaciated peak in the center is Pan de Azucar (4 755 m), the peak to the right is El Campanario, and the valley represented is called Las Lagunillas. Tree in foreground is Gynoxys paramuna. (Photo Cuatrec. C-1219A taken in 1938.)
(Avila and the Silla de Caracas areas) in the Cordillera Costal of north-central Venezuela, Pico de la Neblina along the Venezuela~Brazil border, scattered "humid puna" areas ofthe eastem slopes ofthe Peruvian Andes, the "yungas" region of northeastem Bolivia, scattered areas in Chile and Argentina, and the Itatiaia area of eastem Brazil. For other opinions about the geographical distribution of páramo, see the following: Brack Egg, 1986b; Braun, 1956; Cleef, 1978, 1981b; Cuatrecasas, 1968; Mann, 1968; Monasterio, 1980b; ONERN, 1976; Ribera et al., 1994; Tosí, 1960; Unzueta Q., 1975; Vareschi, 1955; Vuilleumier & Monasterio, 1986.
Clima te The páramos of Colombia and northern Ecuador are influenced by the intertropical convergence of air masses (a low-pressure trough) because oftheir geo-
graphicallocation near the equator. They are generally hum id throughout all or most months ofthe year, with continuous moisture in the form of rain, clouds, and fog, mostly dueto orographic uplift caused by the Andes. Many páramos receive more than 2000 mm of rain annually on their exposed slopes (absolute range 500 toca. 3000 mm/yr). They have a high relative humidity averaging 70~85% (absolute range 25~ 100%). This contrasts with páramos in the northernmost Andes ofVenezuela, the Sierra N evada de Santa Marta in northern Colombia, and in Costa Rica, where there is a marked dry season due to the influence of the northeast Trade winds (Herrmann, 1970, 1971; Lauer, 1979a, 1979b). Páramo becomes driest near its southern limits in southern Ecuador and northern Peru, where they are influenced by two air masses: one from the Amazon basin, which has already re1eased its moisture on the eastern slopes, and another dry cool air mass from the west under the influence of the Humboldt Current. In addition to the large-scale cli-
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MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
matic parameters given above, local microclimates may strongly influence regional weather patterns (Sarmiento, 1986). In northern Peru, the páramo environment grades almost imperceptibly into the "puna" ecosystem, which is more characteristic ofthe central and southern Andean altiplano highlands of central Peru, Bolivia, and Argentina. Puna, in contrast to páramo, is typically xeric and has lower humidity, a shorter wet season, and six months or more of almost no rain. Figure 5 illustrates the overall climate of páramo by the use of climate diagrams from scattered localities. For additional climate diagrams, see Cañadas Cruz, 1983; Cleef, 198lb; Guhl, 198~; J0rgensen & Ulloa U., 1994; Monasterio & Reyes, 1980; Van derHammen et al., 1995b; and Witte, 1994. For comparisons ofpáramo and puna, see reviews in Cabrera, 1957, 1968; Lauer, 1952; QuintanillaP., 1983b;. Sarmiento, 1986; Troll, 1968b; and Young et al., 1997. Páramos have a generally cold and·humid climate with sudden changes in the weather anda diumál fluctuation in temperature from below freezing toas much as 30°C, which often results in a daily freeze-and-thaw cycle that has been referred to as "summer every day and winter every night" (Hedberg, 1964). During the dry season, for example, Páramo Piedras Blancas (Venezuela), at 3700-4700 m elevation, shows exc tremes in air temperatures ranging from -5°C to -11 oc at night to 25-30°C during the day, with freezing temperatures occurring 325-350 nights ofthe year (Pérez, 1987a, 1996b; Pfitsch, 1994). Although overall mean annual temperatures of páramo range from 2°C to l0°C, there is much greater contrast in the climate of higher-elevation areas than is found in lower zones ofthe same mountain ranges. Therefore, the environment becomes harsher and more severe for plant life as altitude increases (Javellas & Thouret, 1995). A typical day often begins cloud-free, cold, crisp, and occasionally windy until mid-moming (Fig. 6); then increased cloud cover from lower elevations, caused by convectional and orographic uplift, brings rain, sleet, fog, and drizzle for much ofthe aftemoon; clearing often occurs in the late afternoon or early evening. Nights are always cold and usually clear with stars filling the ski es; however, frost is frequent in the high páramos and snow is common at the highest altitudes (Fig. 7). During any given day ofthe year, rain, ice, snow, and fog may alternate abruptly with clear sunny ski es and elevated temperatures; in one moment the wet cold necessitates heavy clothing, raincoats, and gloves, while in the next moment, lotion is needed to protect against sunburn. For other general references that discuss páramo climate, see Cuatrecasas, 1968; Eidt, 1968; Guhl, 1974; Lauer, 1981; Monasterio, 1980a; Rundel, 1994;
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Schnetter et al., 1976; Snow, 1976; Sturm, 1978; Sturm & Rangel Ch., 1985; Troll, 1968b; Van der Hammen & Ruiz, 1984; Van der Hammen & Santos, 1995, in press; Van der Hammen et al., 1983, 1989, 1995b; Weber, 1958; and Witte, 1994.
Soils The geology ofthe Andes is extremely variable and consequently so are the soils. Most páramo soils are relatively young and only slightly developed, and are broadly classified into the orders Andosols, Inceptisols, Histosols, Entisols, and Mollisols (Buol et al., 1980; Buringh, 1979; FAO, 1975; Soil Survey Staff, 1975). · · Andosols and Inceptisols include older soil names such as Ando soils, brown tropical soils, black soils, Onji soils, humic allophane soils, hydro humic latosols, volcarric soils, and volcanic ash soils. They are soils formed from and associated with volcanic ash, have a low supply of ( or have lost) bases or iron and aluminum, and show moderate weathering. In páramo, these m ay include Andepts and some Aquepts, as well as Tropepts and Umbrepts. The Andosols most likely to form in páramos are Aquands, Cryands, and Udands. Histosols include older soil names such as bog soils, muck soils, organic soils, and peat soils. They are soils that are highly organic and are found in very wet places such as bogs and swamps. These may inelude Fibrists, Folists, Hemists, and Saprists. Entisols are soils that have little or no evidence of development (i.e., absence of horizons) and have a highly mineral nature. These are often found near snowline and may include Aquents, Fluvents, Orthents, and Psamments. Mollisols are a less common soil group in the páramo, but are very dark colored and base-rich. The most likely group to develop are the Aquolls. At its highest elevations (i.e., superpáramo), páramo soils are very shallowand coarse with a high percentage of rock and sand, there is little to no production of organic matter, and consequently, low water retention (Fig. 8). Superpáramo soils are extremely infertile, for without organic matter or fine grains they have practically no ability to hold exchangeable cations (Pérez, 1992c). Furthern10re, in the superpáramo, the soil surface is recurrently disturbed by needle-ice activity (a type of ground frost), and soilmoving phenomena such as frost-heaving and thawing and sorting of material are common (Pérez, 1987c). In this part ofthe high páramo, the mean annual air temperatures are always low ( 4-6°C), but the cold does not penetra te very deeply into the soil. Soil temperatures, at about 30 cm, more or less reflect those of the mean annual temperatures of the air (Lauer, 1979b). Soil temperature does, however, have a pro-
1999]
PARAMOS
Cerro de la Muerte (3365 m)
7.6" 246r
5
Lo Aguada (3452m)
7.1° 1811
[4]
[19-21]
Páramo lo Culata (3090m)
mm
•e
140
10.13" 1121
mm
[16]
•e
140
IZO
100
50
lOO
40
80
40
80
30
60
60
20
40
40
10
20
20
B.
J
F M A M J
J A S
Lag U!JOS CHingazo {3250 m)
o
5.
J F M A M J
J A S
O N
9.3° 2139
mm
340
969
8.3° 720
Páramo de Berlin (3230m)
mm 120
[ 9]
•e
C,
N D
•e Mucubaji (3550m)
IZO
50
mm
[ 2-1]
160
•e
50
100
40
80
30
60
140 IZO
1~
zo 10
D.
lh_./
40
zo
50
lOO
40
80
30
60 40
JFMAMJJASONO
zo.
F. Laguna del Otúo
50
lOO
40
80
Pampa Galeros (4000m) [ 4-4]
30
60
zo
40
10
zo
G.
•e
o
N
o
4.8°
822
mm 300 280
J
F M A M J
J
A
E.
S O N O
Laguna de Plsoyomtm (3615m)
•e a•
Catopoxl (3560m)
•e
J A S
mm IZO
[1]
•e
J F M A M J
5.5" 929
(4000m)
1150
7.1°
1279
[ 10] 180 160
mm
[10]
140 120
50
100
40
80
30 20 10
H.
~ JFMAMJJ
60 40
zo
ASOND
Fig. 5. C1imate diagrams for páramo (A-l) and puna (J). Walter climate diagrams show the curves ofthe average month1y a ir temperature ("C) and precipitation (mm rainfall) [ 10°C on the ordinate corresponds to 20 mm; hum id periods are shown by vertical! y striped are as; months with greater than 100 mm/month are indicated with black]. Ea eh station' s name, e1evation (m), average monthly temperature (0 C), average year1y precipitation (mm), and number of years of observation [in square brackets] are given. (Sources: A from Stein & Weberling, 1992; B, C from Monasterio & Reyes, 1980; D from Azócar 8F Monasterio, 1979; E, F, I, J from C1eef, 1981 b; G from Witte, 1995b; H from J0rgensen & Ulloa U., 1994.)
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Fig. 6. Páramo de Zipaquirá (Cundinamarca, Colombia) at ca. 3100 abo ve the Valle de Riofiio. Gentle, but consistent winds blow across the top ofthis páramo causing the inflorescences of Espeletia grandiflora to bend. Other associated plants include Puya cryptantha, P. nitida, Paepalanthus columbiensis, and Gaylussacia buxifolia. (Photo Cuatrec. C-1664 taken in 1940.)
found effect on nutrient and water availability, root growth, seed survival and germination, and vegetation zonation (Diemer, 1996; Lauer, 1981; Pérez, 1987b; Smith, 1976; Walter & Medina, 1969a, 1969b). At its middle elevations (i.e., grass páramo), páramo soils are relatively deep, humic, black or dark brownish, and acidic with pH ranging from about 3. 7 to 5.5. They are continuously moist, ore ven saturated with water, dueto the daily formation of dew or frost and the water-retaining capabilities ofthe highly organic, peat-like content. In the lowermost part of the páramo (i.e., subpáramo), near Bogotá (Colombia) and elsewhere in the northem Andes, Sturm ( 1978) has stated that soils share the following properties: dar k color ("black coloured"), moderate to high pH and correspondingly low Ca levels, low free P content, relatively higl). K and N content and reduced uptake of these elements by plants, greater than 10% organic content in the top !ayer, little or no "podzolic" features, and high water capacity. For a discussion of the factors that help to form páramo soils, see Cortés Lombana, 1982, 1995. For
soil types in local páramo studies, see Baruch, 1979; Botero, 1985; Hofstede, 1995c; Pérez, 1991c, 1992c, 1996b; Quintanilla P., 1983b; Rangel Ch., 1989; Salomons, 1986; Sánchez M. et al., 1989; Sevink, 1984; Sturm & Rangel Ch., 1985; Thouret & Faivre, 1989; and Vis in Van der Hammen & dos Santos, 1995. F or other general references about páramo soils, see also Del Llano, 1990; Jenny, 1948; Jenny et al., 1948; Simonson, 1979; Stum1, 1994a; VanderHammen & Ruiz, 1984; Van der Hammen & Santos, 1995, in press; Van der Hammen et al., 1983, 1989; Vareschi, 1970; Wright & Bennema, 1965; and Zéittle, 1970.
Paleohistory and Paleoecology Reconstruction of the paleoecology of páramo and high-elevation montane forest has been the subject of study by Thomas Van der Hammen and his associates since the 1960s. The following is a brief summary of an article by Van der Hammen and C!eef (1986) that emphasizes the paleohistoric events that gave rise to and further influenced development ofthe páramo
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7
Fig. 7. Campsite on the Nevado del Tolima (Tolima, Colombia) at 3800-4200 m. Ovemight snow in a grass páramo with Espe/etia sp., the leaves ofwhich were undoubtedly used for cushion and insulation on the ground. José Cuatrecasas on the left with guides Conne Marulanda, Marco Tulio, and dog. (Photo from glass negative Cuatrec. C-796 taken in 1932.)
ecosystem we see today in the high plain of Bogotá (Colombia). The sequences may be similar or different in other parts of the northern Andes. Unfortunately, detailed accounts from other areas are few in number or not yet available. The Andes began to arise during the Paleocene, and during the Miocene they were probably on the arder of ridges to low mountains up to ca. 1000 m elevation. It was not until the beginning of the Pliocene or slightly later that the northern Andean region uplifted to its present altitudes. During the PlioPleistocene, ca. 4-5 Ma (mi Ilion years ago) there was abundant volcanic activity, at which time elevations above the present treeline carne into being. There may not have been forest y el at the high elevations, since considerable time was needed for the forest line to have risen from around 2500 m (the leve! at that time) up to around 3500 m. The upper Andean forest and páramo belts evolved more or less simultaneously during the Late Pliocene or Early Pleistocene (2-4
Ma). There is evidence that an early páramo vegetation, what Van der Hammen and Cleef (1986) ca11 "protopáramo" vegetation, was present and consisted of Poaceae, Cyperaceae, Asteraceae, Ericaceae, a Polylepis-Acaena (Rosaceae) type of po11en, and Symplocos (Symplocaceae ), Myrica (Myricaceae ), Aragoa (Scrophulariaceae), Hypericum (Clusiaceae), Miconia (Melastomataceae ), Ilex (Aquifoliaceae ), Hydrocotyle (Apiaceae), Borreria (Rubiaceae), Ludwigia (Onagraceae), Polygonum (Polygonaceae ), Valeriana (Valerianaceae), Plantago (Plantaginaceae ), Ranunculus (Ranunculaceae), Myriophyllum (Haloragaceae ), Jamesonia (Pteridaceae ), and Hymenophyllum (Hymenophy1laceae). By 1 Ma (for the high plain ofBogotá, Colombia, at about 2600 111), there is evidence of about 15 to 20 repeated alternations of forest and páramo (i.e., fluctuations of climatic zones) in interglacial and glacial periods. During this time, genera of north temperate origin-such as Alnus (Betulaceae) ca. 1 Ma and Quercus (Fagaceae) ca. 0.3
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Fig. 8. Páramo communities. A. Páramo de Piedras Blancas (Mérida, Venezuela), Mifafí Valley; Pico Los Nevados in the background reaches 4685 m. The upper altitudinal limit of the dense woodlands of Coespeletia lutescens extends into the superpáramo between 4395 m and 4500 m and altemates with largely bare gravelly sand zones clearly differentiated by the sharp contras! in soil color. (Photo by F. L. Pérez taken in 1980.) B. Páramo del Almorzadero (Santander, Colombia) at ca. 3800 m. The superpáramo scree slopes are colonized by severa] species of Senecio, including S. niveo-aureus (see also Fig. 13D), S. almorzaderonis, S. canescens, and S.folidentatus, all characterized by the light-colored, woolly pubescence over much ofthe plant. Other plants often found on superpáramo scree slopes include Werneria pygmaea, Hypochaeris se/osa, Geranium multiceps, Bartsia santo/inifolia, Lachemilla spp., and Plantago monticola. (From J. Cuatrecasas, "Aspectos de la vegetación natural de Colombia," Revista Acad. Colombiana Ci. Exact. 10(40). Reproduced with permission. Photo Cuatrec. C-1974 taken in 1941.) C, D. Desert páramo or "Arenal del Chimborazo" found on the northwest side ofVo1cán Chimborazo (Ecuador) at ca. 3800-4400 m. Strong winds are common in this sand dune type of vegetation dominated by the grasses Stipa hans-meyeri, Calamagrostis mollis, C. ligulata, Cortaderia sericantha, andAgrostis spp., and including scattered p1ants of Bidens hu milis, Loricaria ilinissae, Lasiocepha/us ovatus, L. lingulatus, Pentacalia teretifolius, Senecio nivalis, Astragalus geminiflorus, Geranium ecuadoriense, and Nototriche spp. (Reproduced with pem1ission of the Department of Systematic Botany, University of Aarhus, Denmark. Photo C taken by L. B. Holm-Nielsen in 1980; photo D taken by B. 0llgaard in 1976.)
Ma-appeared in the pollen record and must have passed over a Panamanian landbridge. During the later part ofthe Quatemary, ca. 44,00021,000 B.P. (years before the present) ofthe Last Glacial stadial, glacial and interglacial periods continued to altemate, with sorne short but severe cold periods. At that time there were numerous changes in the proportions between páramo and forest elements, although the páramo flora became well established and dominated the scene. Between ca. 45,000 B.P. and 25,000 B.P., there was a cold and wet period during which glaciation reached its maximum advance. Van der Hammen and Cleef (1986) further explain that during this time the glaciers and forest may even have
been in contact at elevations between 2200 m and 2700 m, and the páramo belt must have been relatively narrow and wet. On the contrary, between 21,000 B.P. and 14,000 B.P. there was a very cold but dry period during which the mean ahnual temperature may have been 6-7°C lower than today. Glaciation was not so extensive, but the páramo belt was broad and dry and covered most ofthe area above 2000 m (i.e., the area where present-day forest occurs). This means that páramo vegetation covered a much greater area than it does today, and that many ofthe currently isolated páramos were then united. It also means that the upper forest line was lowered by 1300-1500 m. At the beginning ofthe Holocene (ca. 10,000 B.P.)
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the climate became much warmer; forest limits rose to elevations even higher than today and páramo vegetation was restricted to above 3300-3500 m. The lower-elevation forest includedDodonaea viscosa (Sapindaceae), Myrica (Myricaceae), Myrsine (= Rapanea) (Myrsinaceae), and Miconia (Melastomataceae); it continued upward withAlnus (Betulaceae) and ended at the highest elevations with Weinmannia (Cunoniaceae) and Quercus (Fagaceae). During the period from 7500 B.P. to ca. 3000 B.P. temperatures rose about 2°C more, causing another upward shift in the forest line of about 300-400 m higher than today and thereby reducing the area occupied by páramo. Finally, at about2900 B.P., there was a noticeab le lowering of the temperature that marked the last downward movement ofthe forest and páramo belts to their present-day positions. Van der Hammen and Cleef (1986) state that the most important changes in the Holocene period have been the temperature changes mentioned above, the development of soils, and the development of peat bogs and soil~ with increased humidity. They summarize by saying that the present-day páramo flora and vegetation is the result of an amalgamation of approximately 4-5 million years of complicated paleohistoric events. Sorne recent publications suggest that the situation may be more complex or may differ in other parts of the Andes than that desc,ribed by Van der Hammen and Cleef (1986). Colinvaux et al. (1997), for example, refute Van der Hammen's idea that Andean vegetation zones were compressed and moved in belts during Quatemary times. Instead, they suggest that during times of glacial cooling or Holocene warming, plant associations showed different spatial diversity and were reformed according to the temperature tolerance of individual species, with heat-intolerant species showing larger displacements. The following list of additional general references about geology, glaciation, and paleohistory in the páramo regions of Central and South America is organized by country: Costa Rica: Hom, 1990b; Weber, 1958. Venezuela: Salgado-Labouriau, 1986; Schubert & Vivas, 1993. Colombia: Helmens, 1990; Hooghiemstra & Ran, 1994; Van der Hammen, 1981b, 1989; Van der Hammen et al., 1973. Ecuador: Colinvaux et al., 1997; Hastenrath, 1981; Wolf, 1892. General Central and South America: Graham, 1973a, 1973b, 1989a, 1989b; Markgraf, 1989.
Vegetation Zonation Over the years, numerous authors have given various names to the different vegetation zones and plant
9
associations within the high Andes. At times the usage of these names can become confusing and it is difficult to know exactly what is being discussed ami/ or compared, but Huber and Riina (1997) and Jorgensen and Ulloa U. (1994) provide summaries of this nomenclature. When thinking of or talking about páramo, for example, reference is generally being made to the open, treeless grasslands with scattered espeletias and shrubs. Much ofthe present-day páramo vegetation of treeless grasslands, however, probably has anthropogenic origins, being maintained by cutting, periodic burning, and grazing-practices that intensified mostly within the last 300 years. Therefore, the natural forest line at which forest ends and undisturbed páramo begins was probably higher than what is seen today, and above 4000 m elevation in sorne places. The evidence for this is that in an undisturbed system there is usually not an abrupt end to the forest, nota sharp edge or border, but instead more of a transition: from the tall forest trees to gradually shorter trees as the elevation increases, then to small trees and shrubs in a more or less thicket formation, and final!y, above the forest line, to the grasses, herbs, and scattered small shrubs ofpáramo (Color Plate III, bottom left). As one reaches the naturallimits of one zone with the next, many ofthe plant species characteristic ofthese zones intermix. It must also be remembered that the boundaries of the vegetation zones and the elevations at which they begin and end are not fixed. Depending on environmental features, such as topography, exposure, soils, and general clima tic conditions, and human intervention, timberline may occur anywhere between about 2800 m and 4000(--4800) m in the northern Andes. Furthermore, on the wetter or windward si de of a mountain or cordillera, the boundary betwe~n forest and páramo (i.e., the contiguous forest line) is higher than on the dry or leeward side (Komer, 1998; L::egaard, 1992; Lauer, 1981, 1993; A. P. Smith, 1975b, 1975c, 1994; Troll, 1958b, 1959, 1968b, 1973; Verweij, 1995). This book follows Cuatrecasas (1934, 1954, 1958, 1968) in dividing páramo ideally into three broad zones based on overall altitude and vegetation structure, with varying degrees of intergradation. From the highest elevations to the lowest, these three zones are called superpáramo, páramo (here referred to as grass páramo), and subpáramo. Superpáramu may be looked upon as the transition belt or ecotone between the permanent snow region above and the grass páramo below. Subpáramo may be seen as the transition belt or ecotone between the grass páramo above and the montane forest below. A brief summary of these three zones follows. Other ideas of páramo zonation or
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MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
modifications ofCuatrecasas's ideas may be found in papers by Monasterio ( 1980c) and Vareschi ( 1970) for Venezuela; by Cleef (1981 b ), Espinal and Montenegro (1963), Fosberg (1944), and Guhl (1982) for Colombia; by Acosta-Solís (1984), Cafiadas Cruz (1983), Harling (1979), J0rgensen and Ulloa U. (1994), and Ramsay (1992) for Ecuador; by Brack Egg (1986b) for Peru; and by Cabrera (1957), Fosberg (1967), and Holdridge (1967) for more general systems of overall vegetation classification that include páramo.
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(Violaceae ); the only páramo gymnosperms, Ephedra americana andE. rupestris (Ephedraceae ); and various species of cryptogams including vagrant ballforming mosses and unattached lichens.
GRASS pÁRAMO
[Cuatrecasas referred to this zone as "grass páramo," "páramo propiamente dicho," or "páramo proper."] At about 3500-4100(-4400) m, vegetation of the grass páramo is continuous and plant cover is generally 100% (Fig. 9; Color Platel). It is composed SUPERPÁRAMO mainly oftussock- or bunch-grasslands dominated by This is a narrow zone of vegetation growing on spe'eies of Calamagrostis and/or Festuca (Color Plates rocky scree and coarse, sandy soils below the snowline I.:_Il). There is a high proportion of dead shoots among at about ( 4000-)4500-4800( -5000) m (Fig. 8). the living that give a yellowish brown or olive-brown Among the three páramo zones it is characterized by to grayish loo k to the grasslands as a whole (Penland, having the lowest air temperature, precipitation, soil · 194la). This is the classic area ofthe genus Espeletia water-holding capacity, and nutrient content, arid by and its relatives (Asteraceae: Espeletiinae), in local communities called "frailejonales," which for so many having the highest solar radiation and night-frost frequency (Baruch, 1984). Plants in this zone must be have cometo symbolize páramo vegetation with their capable of enduring the daily extreme conditions of columnar, woolly, rosette-plant growth form (Color coldness and strong radiation, and regular or frequent Plate Il, bottom left and right). During October and snowfalls. Superpáramo is the zone of least distur- November, in the higher Venezuelan páramos, there bance by humans. It is very localized because of its is no greater display ofbrilliant floral colors anywhere scattered occurrence only on the highest mountains, than when Espeletia schultzii (deep yellow) and Seneand thus has very high endemism. Sorne superpáramos cioformosus (violet-maroon) (Asteraceae), Castilleja occur on mountaintops that are high enough to have fissifolia (bright red, yellow, and green) (Scrophulariglaciers in their uppermost regions, while others are aceae), and Chaetolepis lindeniana (intense magenta) without permanent snow. Vegetation referred to as (Melastomataceae) are in full bloom (Color Plates III, "desert páramo," or sometimes locally called top left, and IV, top left). "arenales," may have so much sand that they loo k like The grass páramo is the most broadly circumbeaches (Fig. SC, D; Color Plate III, bottom right). scribed ofthe three páramo zones. It includes not only At first glance, superpáramo often looks from a the dominant grassland communities but also the distance to be bare ground. In reality, however, it is greatest number of azonal communities, which are home to tiny, clumped or scattered plants such as determined by specific factors such as soil moisture Azorella pedunculata (Apiaceae); Hypochaeris and topography, and páramo growth forms. The most sessiliflora, Senecio canescens, S. nivalis, S. frequently encountered azonal communities are deadglacialis, S. supremus, S. comosus, Pentacalia scribed below under the grass páramo, but severa! are gelida, andXenophyllwn rigidum (= Werneria rigida) also found to a lirnited extent in the superpáramo and (Asteraceae); Draba pachythyrsa, D. depressa, and subpáramo zones as well. The grassland comrnunities Eudema nubigena (Brassicaceae); Arenaria spp. and of the grass páramo zone have suffered greatly frorn Cerastium jloccosum (Caryophyllaceae ); Pernettya burning and grazing (see "Impact of Burning and pros trata and Disterigma empetrifolium (Ericaceae ); Grazing," below). Astragalus geminifloms, Lupinus alopecuroides, L. Grass páramo rnay consist oftall- and short-grass microphyllus, and L. smithianus (Fabaceae); Gera- communities ("pajonales," "pastizales," or "prados") nium multipartitum (Geraniaceae); Luzula racemosa that include both herbaceous and woody vegetation, (Juncaceae); Nototrichejamesonii and N. chimbo- but they are dominated by tussock or bunchgrasses razoensis (Malvaceae ); Aciachne pulvinata, Agrostis (Fig. 1OA, B; Color Plate I, bottom right). In the tallnigritella, Bromus oliganthus, Poa cucullata, P. grass communities, with grass to 1 m tall, Calamatrachyphylla, P. orthophylla, and Stipa ichu (Poaceae); grostis recta usually dominates on drier sites and C. Plantago sericea subsp. nubigena (Plantaginaceae); effusa dominates on moister. Short-grass communiValeriana alpifolia (Valerianaceae); Viola pygmaea ties usually indicate grazing and burning pressures and
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Fig. 9. Páramo communities. A. Climbing Volcán Puracé (Cauca, Colombia) at ca. 4200 m. The dominants in this rocky-soil vegetation are the clump-forming grassFestuca tolucensis and the dark-colored shrub Loricaria thuyoides. Other associated plants include Agrostisfoliata, Luzula gigantea, and Lasiocephalus puracensis. (From J. Cuatrecasas, "Páramo vegetation and its Ji fe forms," Col!. Geogr. 9. Reproduced with permission. Photo Cuatrec. C-2103 taken in 1943.) B. Climbing Nevado del Cocuy above Pozo Azul (Boyacá, Colombia) at ca. 4150 m. In the foreground an open hillside is dominated by the grass Calamagrostis ,e./fusa and the asteraceous Espeletiopsis colombiana. Other associates in this community include Oritrophium peruvianum, Baccharis tricuneata, Hypericum prostratum, H. mexicanuin, Aciachne sp., Jamesonia sp., Bartsia laniflora, Castilleja sp., and Geranium sp. (Photo Cuatrec. 1225A taken in 1938.) C. Grass páramo with Espeletia sp. below the Nevado del Tolima (Quindío, Colombia) at ca. 4000 m. (Photo Cuatrec. C-803 taken in 1932.) D. Páramo de Cruz Verde abo ve Santafé de Bogotá (Cundinamarca, Colombia) at ca. 3400 m. Páramo with shrubs of Diplostephium heterophyllum, Pentacaliaflos~fragrans, P. vaccinioides, Senecio subruncinnatus, Hypericum laricifolium, H. thuyoides, Berberís sp., and Aragoa abietina, rosettes of Puya trianae and P. santosii, the bamboo Chusquea tesse/lata and bunchgrasses of Calamagrostis e.lfusa and Festuca dolichophy/la, and herbs such as Halenia adpressa, Gentianella spp., Carex pichinchensis, Valeriana pilosa, Paepalanthus karstenii, P. columbiensis, Geranium multiceps, and Bartsia santolinifolia. (From J. Cuatrecasas, "Páramo vegetation and its life forms," Col!. Geogr. 9. Reproduced with permission. Photo Cuatrec. C-1740 taken in 1940.)
are often dominated by species of Agrostis, Fes tuca, and Paspalum. The grass Aciachne pulvinata often forms low or flat cushions with very sharp (to the touch) leaves in drier-site short-grass páramos. In Colombia, dwarfbamboos (Chusquea spp.) domínate on slopes with very wet clima tes in communities known as bamboo brakes or "chuscales." If one takes the time to search between the bunchgrasses of the open grass páramo, there is an astonishing array of species of small herbs: Bomarea spp. (Alstroemeriaceae) (Fig. !lA); Eryngium humile
(Apiaceae); Perezia spp. (Asteraceae); Lysipomia spp. (Campanulaceae-Lobelioideae); Paepalanthus (Eriocaulaceae) (Fig. 12A); Lupinus spp. (Fabaceae) (Fig. 12C; Color Plate VI, bottom left), Gentiana sedifolia, Gentianella spp. (Color Plate IV, top right), and Halenia spp. (Gentianaceae); Lachemilla spp. (Rosaceae) (Figs. 12B, l3A); Gunnera magellanica (Haloragaceae) (Fig. l3C); Sisyrinchium spp. and Orthrosanthus chimboracensis (Iridaceae) (Color Plate VI, top right); Oenothera epilobiifolia (Onagraceae); Ranunculus spp. (Ranunculaceae);
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Fig. 10. Páramo communities. A. Páramo Almorzadero (Santander, Colombia) at ca. 3800 rrt. Wet, short-grass páramo ("prado turboso pantanoso") with Espeletia estanislana and the grass Calamagrostis bogotensis. Other associates include Espeletia conglomerata, Diplostephium revolutum, Hypericum sp., and Castratella piloseloides. (From J. Cuatrecasas, "Aspectos de la vegetación natural de Colombia," Revista Acad. Colombiana Ci. Exact. 10(40). Reproduced with permission. Photo Cuatrec. C-1705 taken in 1940.) B. Macizo de Bogotá, Quebrada de San Cristobal, abo ve El Delirio, near Santafé de Bogotá (Cundinamarca, Colombia) at ca. 2950 m. Heavily disturbed, short-grass vegetation ("prado") of Lachemilla orbiculata with Hypochaeris sessiliflora (see close-up photo in Fig. 13A) andAa rostrata, in a disturbed shrub community ("matorral") consisting of Viburnum triphyllum, Tibouchina grossa, Perseo mutisii. (Photo Cuatrec. C-1599 taken in 1940.) C. Páramo del Escoba! on ascent to Nevado del Cocuy (Boyacá, Colombia) at ca. 3700 m. Residual forest with shrubs and small trees of Polylepis sp. (note peeling bark on small tree to right), Esca/lonia myrtilloides, Gynoxys sp., Miconia salicifolia ( on left), Espeletiopsis jimenez-quesadae (center right), Espeletia lopezii (lower plants to left), Hypericum mexicanum, Ageratina sp., and Baccharis sp. Other herbaceous plants include Acaena sp., Gentiane/la sp., Sisyrinchium sp., and Orthrosanthus chimboracensis. (Photo Cuatrec. C-1244 taken in 1938.) D. Ascent ofNevado del Cocuy near La Cueva (Boyacá, Colombia) at ca. 3700 m. Shrub páramo with small tree of Esca/lonia myrtilloides (center) parasitized by Aetanthus dichotomus (see close-up photo in Fig. 14B). On the left a shrub of Miconia salicifolia. Other plants include Espeletiopsis jimenez-quesadae, Orthrosanthus chimboracensis, andAcaena cylindrostachya. (Photo Cuatrec. C-1204A taken in 1938.)
Castillejo fissifolia and Bartsia spp. (Scrophulariaceae); the lycopods Huperzía spp.; and the fem genus Jamesonía (Pteridacae) (Color Plate IV, bottom right). Common species rooted within the tussock formations include Cerastium spp. (Caryophyllaceae), Vicia andicola (Fabaceae), Geranium spp. (Geraniaceae) (Fig. 13D), and Bromus lanatus (Poaceae). Rumex acetosella (Polygonaceae) is a weedy adventive species that often forms a red ground cover over large disturbed areas, usually after potato or wheat cultivation (Color Plate III, top right). Senecío níveo-
aureus (Asteraceae) (Fig. 12D) and related species in Senecío sect. Culcitíum (Fig. llB) add a stark beauty
with their dense, woolly, whitish to pale yellowish pubescence, which covers practically the entire plant. Also common in the grass páramo are acaulescent rosette-plants in the genera Oreomyrrhis (Apiaceae), Hypochaeris (Asteraceae) (Fig. 13A), Acaulímalva (Malvaceae), and Acaena (Rosaceae), and cushionplants in genera such as Wernería and Xenophyllum (Asteraceae) (Color Plate VI, top left), Draba (Brassicaceae ), Arenaría (Caryophyllaceae), andPaepalanthus
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Fig. 11. Páramo herbs. A. Bomarea pauciflora from area of San Miguel, SW ofthe Sabana de Bogotá (Cundinamarca, Colombia) at ca. 2900 m. Flowers orange when fresh. (Photo Cuatrec. C-1858 taken in 1941.) B. Senecio santanderensis from Páramo Almorzadero (Santander, Colombia) at ca. 3800 m. Note dense white, woolly pubescence covering plant. (Photo Cuatrec. C1976 taken in 1941.) C. Arcytophyllum nitidum from area ofQuebrada de Chico near Santafé de Bogotá (Cundinamarca, Colombia) at ca. 2650 m. Flowers white when fresh. (Photo Cuatrec. C-1393 taken in 1939.) D. Chuquiragajussieui ("cartagena") from páramo along the Quebrada de Las Vegas (source ofthe Río Tuluá) (Valle, Colombia) al ca. 3500 m. Capitula with involucra] bracts bright orange-yellow when fresh and the styles vivid red; the leaves are brilliant green. (Photo Cuatrec. C-2397 taken in 1946.)
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Fig.12. Páramo herbs. A. Paepalanthus sp. from páramo on the Macizo de Bogotá, region ofGuadalupe, above Santafé de Bogotá (Cundinamarca, Colombia) at ca. 3220 m. Plant association includes Espe/etia grandiflora, Puya goudotiana, and Ca/amagrostis sp. (Photo Cuatrecasas C-1392 taken in 1939.) B. Along the Quebrada de Las Vegas (source ofthe Río Tuluá) (Valle, Colombia) at ca. 3500 m. Wet páramo meadow with a mixture ofIsoetes killipii, Plagiochei/us solivaeformis, Lachemil/a niva/is, Sisyrinchium sp., Ranunculus spp., Caro: bonplandii, Jso/epis inundata, and Oreobolus obtusangu/us. (From J. Cuatrecasas, "Aspectos de la vegetación natural de Colombia," Revista Acad. Colombiana Ci. Exact. 10(40). Reproduced with permission. Photo Cuatrec. C-2398 taken in 1946.) C. Lupinus alopecuroides, an acaulescent rosette plant, from moist bunchgrass area of Páramo del Puracé (Cauca, Colombia) at ca. 3900 m. Note columnar inflorescence with paJe blue flowers (when fresh). (From J. Cuatrecasas, "Aspectos de la vegetación natural de Colombia," Revista Acad. Colombiana Ci. Exact. 10(40). Reproduced with permission. Photo Cuatrec. C-21 06 taken in 1943.) D. Senecio niveo-aureus from short-grass area ofPáramo del Escoba! (Boyacá, Colombia) around La Cueva at ca. 3800 m. Plants are ca. 1 m tall and densely white villose; the ray flowers are yellow-gold when fresh. Other plants in this community included Senecio formosoides, Carex bonplandii, Jsolepis inundata, Gunnera mage/lanica, Geranium stramineum, Ha/eniafo/iosa, and Jamesonia cinnamomea. (Photo Cuatrec. C-1212A taken in 1938.)
(Eriocaulaceae). Grass páramo is also rich in small and large shrubs, such as those genera mentioned below in the section on subpáramo, buÚhey are more scattered in the grass páramo (Color Plate II, top left). They include many species of Baccharis, Diplostephium (Fig. 14A), Gynoxys, Loricaria (Color Plate V, top left), and Pentacalia (Asteraceae ); Hypericum (Clusiaceae); Gaylussacia, Gaultheria, Pernettya, and Vaccinium (Ericaceae); Arcytophyllum (Rubiaceae) (Fig. llC); and Valériana (Valerianaceae). Swampy or boggy azonal sites, called "cushion mires" or "turberas," are common, especially in the uppermost grass páramo (Bosman et al., 1993; Cleef,
1980a; Fig. 15A; Color Plate II, top right). Here species ofthe spectacular cushion-plant growth form attain their best development, e.g., Azorella aretioides, A. multzfida, and A. pedunculata (Apiaceae); Oreobolus obtusangulus (Cyperaceae ); the moss-like Distichia musco ides (Juncaceae ); andP!antago rígida (Plantaginaceae). These long-lived cushions often form the substrate for other, smaller plants (as epiphytes) such as Hypochaeris spp. (Asteraceae), Carex spp. (Cyperaceae), Disterigma empetrifolium and Pernettya pros trata (Ericaceae ), Gen tia na sedifolia and Gentianella spp. (Gentianaceae), Agrostis spp. and Poa spp. (Poaceae), and Lachemilla spp. (Rosaceae). These cushion "epiphytes" derive their wa-
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Fig. 13. Páramo herbs. A. Lachemilla orbiculata, the plants with rounded 1eaves and crenate margins, but without flowers and Hypochaeris sessi/iflora, the plant with yellowish-orange (when fresh), sessile ray flowers, forming a thick mat in a heavily disturbed, short-grass ("prado") vegetation on the Macizo de Bogotá along the Quebrada de San Cristobal near Santafé de Bogotá (Cundinamarca, Colombia) at ca. 2950 m (see Fig. 10B). (Photo Cuatrec. C-1598 taken in 1940.) B. Castratella pi/loseloides from the Macizo de Bogotá between La Viga and Diego largo (Cundinamarca, Colombia) at ca. 3380 m. This endemic, rosette-forming Me1astomataceae (with yellow flowers when fresh) grows in the moist, shortgrass páramo community along withAzorel/a sp., Valeriana pilosa, Lachemi/la sp., Niphogeton glaucescens, Calamagrostis effusa, Huperzia hohenackeri, L. c/avatum, and the shrubs Miconia buxifolia, Arcytophyllum muticum, Ugni myricoides, Hypericum spp., and Diplostephium phylicoides (From J. Cuatrecasas, "Aspectos de la vegetación natural de Colombia," Revista Acad. Colombiana Ci. Exact. 1O(40). Reproduced with permission. Photo Cuatrec. C-1405 taken in 193 9.) C. Gunnera megallanica from wet páramo meadow along the Quebrade de Las Vegas (source of Río Tuluá) (Valle, Colombia) at ca. 3500 m (see a1so Fig. 12B). (Photo Cuatrec. C-2396 taken in 1946.) D. Geranium multiceps, the plant with pinkish-white flowers (when fresh) with the bunchgrass Calamagrostis effusa from Páramo de Cruz Verde above Santafé de Bogotá (Cundinamarca, Colombia). See Fig. 9D for associated species in this community. (Photo Cuatrec. C-1738 taken in 1940.)
ter and nutrients from the process of litter decomposition and nutrient release taking place within the cushion itself(Sklenáf, 1998). Other common species of swampy or boggy sites include Juncus spp. (Juncaceae ), the páramo endemic Castratella piloselloides (Melastomataceae) (Fig. 13B), and Valeriana spp. (Valerianacee). Also sometimes abundant are the lichen genera Cladia (Cladoniaceae) and Usnea (Parmeliaceae); mosses Campylopus (Dicranaceae), Breutelia (Bartramiaceae), and Sphagnum (Sphagnaceae); and liverworts Riccardia (Aneuraceae), Frul-
lanía (Jubulaceae), andLophozia (Jungermanniaceae). Other wet or flooded azonal communities such as marshes ("pantanos" or "ciénagas"), seeps, and springs (Fig. 15B) may also include bunchgrasses (especially Festuca dolichophylla), but with more species of sedges ( Carex, Eleocharis, etc.) and mosses such as Drepanocladus (Amblystegiaceae) and Sphagnum (Sphagnaceae). Also found are the various species of Lilaeopsis and Hydrocotyle (Apiaceae), Oritrophium peruvianum (Asteraceae ), Plagioboth1ys linifolius (Boraginaceae), Draba lindenii (Brassi-
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Fig.14. Páramo shrubs. A. Diplostephium phylicoides from area oiSan Miguel, SW ofthe Sabana de Bogotá (Cundinamarca, Colombia) at 2900 m. Ligulate flowers blue when fresh. (Photo Cuatrec. C-1853A taken in 1941.) B. Aetanthus dichotomus from aseen! ofNevado del Cocuy near La Cueva (Boyacá, Colombia) at ca. 3700 m. This plant is a parasite on small tree of Escal/onia myrtilloides in a shrub páramo (see Fig. lOD formore associates in this community). Flowers have red corollas with yellowthroat and lobes when fresh. (Photo Cuatrec. C-1828 taken in 1941.) C. Aragoa abietina from the Macizo de Bogotá, Quebrada de Chico area near Santafé de Bogotá (Cundinamarca, Colombia) at 2800---3000 m. It was associated with Gay/ussacia buxifo/ia, Hypericum sp., Dip/ostephium ochraceum, and Espe/etia grandij/ora. (Photo Cuatrec. C-1404 Ú1ken in 1939.) D. Berberís cuatrecasasii ("grano de oro") from Páramo de Bavaya (= Paramillo de Santa Lucía) near Corrales in the Cuchilla de Barragán (Valle, Colombia) at 3500 m. Part of a shrub community consisting of Hypericum laricifolia, Hesperomeles sp., Bejaria spp., Gynoxys laura/a, Tibouchina grasa, Tristerix sp., Puya sp., and Valeriana sp. (Photo Cuatrec. C-2391 taken in 1946.)
caceae ), Stellaria media (Caryophyllaceae ), Halenia spp. (Gentianaceae), Myriophyllum quítense (Haloragaceae), Juncus spp. (Juncaceae), Huperzia spp. (Lycopodiacee), Rumex tolimensis (Polygonaceae) to 4-5 m tall, Caltha sagittata and Ranunculus praemorsus (Ranunculaceae), andMimulus glabratus and Pedicularis in curva (Scrophulariaceae ). Shallow pool, lake, and river communities (Fig. 15C, D) inelude Callitriche spp. (Callitrichaceae), E latine spp. (Elatinaceae ), Myriophyllum spp. (Haloragaceae ), Isoetes spp. (Isoetaceae), Cortaderia spp. (Poaceae), Potamogeton spp. (Potamogetonaceae), and Ranunculus spp. (Ranunculaceae). Roe k ledge and cliff communities harbar another distinct group of interesting plants including Draba
spp. (Brassicaceae ), Luzula racemosa (Cyperaceae), Escalla nía myrtilloides and Ribes hirtum (Grossulariaceae), Calceolaria spp. (Scrophulariaceae) (Color Plate IV, bottom left), and numerous fem species in the generaAsplenium and Cystopteris (Aspleniaceae) and Elaphoglossum and Woodsia (Dryopteridaceae). Trees ofthe genus Polylepis (Rosaceae), with their characteristic reddish, exfoliating bark and strangely contorted trunks and branches, may form localized, isolated woodlands or forest communities to over 4000 m elevation within otherwise grass páramo (Fig .. !OC). They are often found on scree slopes, near the shelter ofrock cliffs, or in river valleys (Color Plate I, bottom left). It is interesting that within the Polylepis forest there is a noticeable drop in plant diversity as
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Fig. 15. Páramo communities. A. Cushion bog or "turbera" dominated by Distichia tolimensis floating on swampy ground in páramo ofthe Nevado del Cocuy (Boyacá, Colombia) at ca. 4200 m. Between the cushions are Calamagrostis sp., Xenophyllum crassa, Plagiocheilus solivaeformis, a few stems of Pernettya prostrata, and Sphagnum spp. (From J. Cuatrecasas, "Páramo vegetation and its life forms," Col l. Geogr. 9. Reproduced with permission. Photo Cuatrec. C-1231 taken in 1938.) B. Marshy area or "pantanoso" along meandering river in the Valle de Las Lagunillas near the Nevado del Cocuy (Boyacá, Colombia) at ca. 3900 m. The wet, short grasslands are dominated by Aciachne sp. with Agrostis trichodes, Sisyrinchium pusillum, and Halenia insignis; the marshy areas contain Espe/etia lopezii, while the slopes are occupied by Espeletiopsis colombiana, Pentacalia vaccinoides, Diplostephium /acunosum, and Baccharis sp. (Photo Cuatrecasas C-1215A taken in 1938.) C. Shallow pool or "lagunita" on the Planos del Toldadero (near locality of Fig. l5B) at ca. 3900 m. The same species as in Fig. 15B domínate the surrounding areas, whereas the pools have aquatic vegetation such as Ranunculus spp., Plagiocheilus solivaeformis, and Isoetes spp. (Photo Cuatrec. C-1235 taken in 1938.) D. Glaciallake in superpáramo immediately below the Nevado del Cocuy (Boyacá, Colombia) at ca. 4500 m. Agrostis nigritella is one ofthe first species to colonize the wet areas between the rocks. (From J. Cuatrecasas, "Aspectos de la vegetación natural de Colombia," Revista Acad. Colombiana Ci. Exact. 10(40). Reproduced with permission. Photo Cuatrec. C-1229 taken in 1938.)
compared to the surrounding grass páramo (páramo species are probably shade intolerant), and furthermore that certain plants grow only under Polylepis. Polylepis forest is still relatively underexplored, and an overall study of its flora would be very interesting (but see Amal, 1983; Fjeldsa & Kessler, 1996; and Hueclc, 1960a). SUBPÁRAMO
The lowest zone, called subpáramo, is also the most diverse and floristically showy. It is a shrub-
dominated transition zone at (2800-)3000-3500 m comprising elernents from forest below and the grass páramo above (Fig. 16). It is often a mosaic of shrubs and small scattered trees, gradually reduced in size, giving way to scrub and low vegetation of dwarf shrubs, grasses, and herbs ofthe grass páramo above (Fig. lOD). Sometimes local changes in topography and soils may give way to different microclimatic conditions, so small patches of forest may be present (Color Plate II, top left). Where the ecotone between forest and páramo is abrupt, or where isolated shrubtree islands are found within páramo, it is usually
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Fig. 16. Páramo communities. A, B. Páramo de Santa Lucía, just south ofthe Laguna de La Cocha (Putumayo, Colombia) at ca. 3100 m. A. Subpáramo woodland withEspeletia sp., Calamagrostis cf. bogotensis, Chusquea tessellata,Blechnum auratum, Weinmannia cf. cochensis, Hypericum lancioides, Diplostephiumjloribundu, Miconia sp., and Clusia multiflora. (Photo Cuatrec. C-1824 taken in 1941.) B. General view ofthe Espeletia woodland or "frailejonal." (Photo Cuatrec. C1827 taken in 1941.) C; Páramo de San Antonio del Bordoncillo (Putumayo, Colombia), between Santiago de Sibundoy andEl Encano, at ca. 3250 m. Subpáramo remnant forest with small trees and shrubs of Weinmannia cochensis, Clusia multiflora, Clethra ovalifolia, Hesperomeles pachyphylla,Myrsine dependens, Oreopanax nigrus, Hedyosmum cumbalense, Gaiadendron punctatum, Miconia nodosa, M ligustrina, Ilex myricoides, Diplostephium hartwegii, D. bicolor, Baccharis granadina, Semiramisia pulcherrima, Themistoclesia epiphytica, Plutarchia angulata, Monnina obtusifolia, Fuchsia petiolaris, the tree fern Blechnum auratum, and rosette species Espeletia pycnophylla and Puya vestita, the grasses Calamagrostis bogotensis, Neurolepis acuminatissima, and Chusquea tessellata, and the sedge Rhynchospora sp. (Photo Cuatrec. C-1819 taken in 1941.) D. Shrub páramo/subpáramo NE ofPamplona (Norte de Santander, Colombia) at ca. 2770 m. In the foreground are acaulescent rosettes of Espeletiopsis santanderensis and behind themEspeletia brassicoidea. In the background is a matorral with small trees of Weinmannia tomentosa, shrubs of Arcytophullum nitidum, Be}aria resinosa, Gaultheria erecta, Plutarchia coronaria, Chaetolepis microphylla, Monochaetum bonplandii, Miconia spp., Hypericum spp., and Ternstroemia meridionalis, and herbaceous rosettes of Paepalanthus columbiensis. (Photo Cuatrec. C-1722 taken in 1940.)
brought about by anthropogenic means, especially cutting, buming, and cattle-grazing. Therefore, it is possible that subpáramo consists ofnearly all secondary communities. The subpáramo communities, often known as scrub, thickets, "chaparrales" or "·mattorales," are composed predominantly of shrubby or woody vegetation that is sometimes lacking or rare in the Andean forest below. These include species ofthe general!ex (Aquifoliaceae), Ageratina, Baccharis, Chuquiraga
(Fig. llD), Diplostephium (Fig. 14A), Gynoxys, Loricaria, Senecio and Stevia (Asteraceae ), Berberís (Berberidaceae) (Fig. 14D), Siphocampylus (Campanulaceae-Lobelioideae), Hypericum (Clusiaceae), Coriaria (Coriariaceae), Desfontainia (Desfontainiaceae), Bejaria, Cavendishia, Gaultheria, Afacleania, Pernettya, Semiramisia (Color Plate Vl, bottom right), Themistoclesia and Vaccinium (Ericaceae), Brachyotum, Chaetolepis (Color Plate III, top left), Miconia and Monochaetum (Melastomataceae), Myrsine
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(Myrsinaceae), Monnina (Polygalaceae ), Rubus (Ro- tion, alliance, order, class, etc.). For further explanasaceae), Arcytophyllum (Rubiaceae) (Fig. l!C), tion ofthis methodology, see Becking, 1957; BraunAragoa (Fig. 14C) and Calceolaria (Scrophulari- Blanquet, 1964; Mueller-Dombois & Ellenberg, 1974; aceae), Symplocos (Symplocaceae ), and Ternstroemia and Westhoff & Van der Maarel, 1973. For applied (Theaceae ). Also common are woody epiphytes in the examples of this methodology, with lists of species Ericaceae ( e.g., Disterigma, Plutarchia, Sphyro- characterizing the various plant communities, see spermum, Thibaudia) and Loranthaceae (e.g., Aetan- Aguirre & Rangel Ch., 1976; Cleef, 198lb (includthus) (Figs. 10D and 14B). ing figs. 11-79); Franco R. et al., 1986; Lozano C. & Genera forming low forests of small trees up to eight Schnetter, 1976; Sturm & Rangel Ch., 1985 (includmeters tal!, often in tree islands known as "bosques ing figs. 4-7); and Vargas Ríos & Zuluaga, 1980. achaparrados," include Oreopanax and Schefflera Páramo studies using other quantitative methods may (Araliaceae), Gynoxys, Diplostephium and Senecio be seen in works by Baruch (1984) and Fariñas and (Asteraceae), Buddleja (Buddlejaceae), Weinmannia Monasterio (1980) for Venezuela, and by Balslev and (Cunoniaceae), Bejaria (Ericaceae), Escallonia de Vries (1982), Grubb et al. (unpubl.), Muñoz et al. (Grossulariaceae), Miconia (Melastomataceae), Myrsine (1985), and Ramsay (1992) for Ecuador. (Myrsinaceae), and H esperomeles (Rosaceae). Subpáramo has many common names, of which Morphological and Physiological Adaptations the most frequently used are páramo forest, páramo thicket, shrubby páramo, subpáramo woodland, Since páramo is a high-elevation tropical ecosyssubpáramo chaparral, subpáramo elfin forest, and tem, certain characteristic physical, chemical, and tropical subalpine forest (in English); paramillo, climatic features affect the biological functioning of paramito, bosquecillo de páramo, matorral de páramo, the organisms that live there. Therefore, the plants that matorral de subpáramo, bosque paramero, bosque grow in these areas must be adapted to the following subparamero, bosque enano, bosque musgoso de conditions: 1) high-elevation air (with less water consubpáramo, bosquete andino, and (in Venezuela) tent and lower partía! pressures of gasses such as 0 2 chirivital (in Spanish); and paramillo thicket and and CO); 2) low temperatures ( the annual mean temperature in the shade decreases about 0.6°C for each paramillo scrub (mixed Spanish and English). In general terms, subpáramo is the most difficult 100 m increase, whereas in full sunlight it increases of the three páramo zones to define, because it has with elevation because the atmosphere neither absorbs been greatly extended and expanded both horizontal! y nor disperses as much radiation energy as in the lower and vertical! y (downwards) by human disturbance and regions); 3) intense ultraviolet radiation (highest in habitat destruction over hundreds, perhaps thousands, equatorial high mountains, but controlled by frequent ofyears. This is due especially to cutting and burning fog and cloud cover); 4) rapid changes in insolation at the upper forest line for herding and agricultura! resulting in quick absorption or loss of heat; 5) drypurposes. Correspondingly, forest lines appear to have ing effect ofwinds (which can increase transpiration been lowered by as much as severa! hundred meters to the point of desiccation); 6) physiological dryness by human interference (Lregaard, 1992), and rpany (due to the combination of low temperatures, intense species that were more restricted to relatively high transpiration during sunny periods, and drying effects elevations appear to have colonized lower life zones ofwinds, along with high soil acidity and high osmotic (Budowski, 1968). Much ofboth subpáramo (noncon- pressure of soil water making root water-absorption troversial) and grass páramo (somewhat controversia!) difficult); and 7) physical damage from hail and posnow occur in areas that in the past were probably cov- sibly snow (Acosta-Solís, 1984; Cleef, 1981 b; Cuatrecasas, 1968; Little, 1981; Mani, 1980; ered with upper montane forests (Uegaard, 1992). Sarmiento, 1986; Smith, 1981; Vareschi, 1970; Detailed classifications ofpáramo vegetation have Young, pers. comm.). Consequently, growth and debeen constructed utilizing relevés in the Zürich- composition are slow, primary productivity is low, and Montpellier approach and described following the natural succession of the vegetation takes a long time, Code of Phytosociological Nomenclature (Barkman especially when woody species are involved et al., 1976). This system divides vegetation into units (Ferweda, 1987; Hofstede, 1995c; Hom, 1989a, 1997; or communities on the basis of floristic, physiogno- Janzen, 1973; Ramsay & Oxley, 1996; Salamanca V., mic, and ecologic similarities, and usual! y employs a 1991; Sarmiento M. et al., 1990; Smith, 1981; Sturm, detailed classification ofthe vegetation consisting of 1978; Williamson et al., 1986). It is important to rea syntaxonomical hierarchy (subassociation, associa- member that there is no strong temperature seasonal-
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MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
ity, no marked change from summer to winter as in the temperate regions; rather, growth in the páramo is continuous throughout the year, and great changes in temperature (and precipitation, toa certairt'extent) occur every da y ( diumally). Smith and Young (1987) noted that "many aspects of morphology and physiology séem to provide escape from, or tolerance to, extreme diurna! climatic fluctuations." Recent studies have shown how giant Andean rosettes (e.g.,Espeletia and Puya) have evolved adaptations that favor temperature insulation and the maintenance of a positive water balance under the severe conditions of the páramo environment. Adaptations to low temperatures include freezing-avoidance mechanisms such as supercooling of adult leaves (Goldstein et aL, 1985b; Larcher, 1975; Rada R. et aL, 1985a); insulation by retention of dead leaves (marcescent leaves), which protects rosette stems from freezing (Goldstein & Meinzer, 1983; Rada R. et aL, 1985a; Smith, 1979); parabolic 1eaf geometry and nyctinastic movements ofthe leaves, which result in protection of the apical leaf bud from freezing (Hedberg, 1964; Larcher, 1975; Smith, 1974b); thermal buffering by mucilaginous fluids secreted by the leaf bases, which protects apical buds (Smith & Y oung, 1987); dense leaf pubescence, which reduce~ transpiration (Baruch, 1972) and increases leaftemperature (Baruch, 1975; Meinzer & Goldstein, 1985; Meinzer et aL, 1994a; Miller, 1986, 1994); tal! aerial stems, which protect buds against the low mínimum nighttime temperatures at ground leve] (Meinzer et aL, 1994a; A. P. Smith, 1980); and contractile roots in juvenile plants, which draw the developing stem into the ground (Smith in Smith & Y oung, 1981). Rosette adaptations to 1ow moisture levels include changes in the method of C0 2 assimilation, such as a switch to the CAM photosynthetic pathway (Baruch & Smith, 1979; Medina, 1974) anda well-developed water-storing pith tissue that can be used particular! y during early moming hours, when cold or frozen soils limit water uptake during that period ofhigh transpiration (Goldstein & Meinzer, 1983; Goldstein et aL 1984; Meinzer & Goldstein, 1986; Meinzer et aL, 1985, 1994b ). Sorne rosette plants, like Draba chionophila (Brassicaceae), which grows at elevations to ca. 4800 m in the Venezuelan Andes, are freezetolerant (i.e., freezing injury occurs only when temperatures drop below the temperature at which extracellular ice formation begins) (Azócar et aL, 1988; Goldstein et aL, 1994; Pfitsch, 1994). For reviews of general adaptive radiation in Espeletia and other plants ofthe high Andes, see also Beck, 1994; Goldstein et aL, 1994; Hedberg & Hedberg, 1979; Monasterio,
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1986b; Monasterio & Sarmiento, 1991; Ramsay, 1992; Rundel et aL, 1994b; and Smith & Young, 1987. Sorne ofthe physiological and morphologi'cal adaptations discussed above work in combination~i.e., not only as avoidance/tolerance mechanisms against a cold environment but also as useful adaptations against a hot environment resu1ting from frequent tires (see Fig. 17). Adaptations to high elevation by páramo p1ants result in the characteristic growth forms discussed below.
Growth Forms Most high-elevation tropical plant communities háve a characteristic physiognomy that repeats in geographically disjunct areas ofthe world where they ·occur~e.g., South America, East Africa, and Hawaii ~Cuatrecasas, 1968; Hedberg, 1964, 1992; Hedberg & Hedberg, 1979; Raunkiaer, 1934; Troll, 1958b; Vareschi, 1970). The growth forms (sensu lato) that characterize this physiognomy are examples of convergent evolution, the forms having evolved independently in severa] different plant families on distant continents in response to the unique high-altitude tropical environments. These growth forms often result from the ecological and morphological adaptations mentioned above. Many are also apparently adapted to survive fire (L::egaard, 1992; Young & León, 1991). While the tree is the dominant growth form in the forest, it is essentially absent from páramo (except the genus Polylepis anda few associated species). Apparently it cannot survive at such elevations. In fact, tree growth ceas es when soil temperatures drop to 6-1 0°C (Larcher, 1975; Lauer, 1979a, 1981; Walter & Medina, 1969a, 1969b). Páramo, however, has its own important and conspicuous growth forms (not al! of which are strictly found in páramo), such as bunchgrasses, rosette plants, cushion plants, microphyllous and dwarf shrubs, vagrant plants, and geophytes. The presence of rosette plants (some of giant size), for example, is probably a good general indicator of páramo and seems to be one of the clearest distinctions between high-elevation areas of tropical and temperate latitudes (Smith, 1994). The most characteristic growth forms of the high-elevation páramos are summarized below. See also Balslev & de Vries, 1991; Hedberg, 1964 (for comparisons with Afroalpine plants); Hedberg & Hedberg, 1979; and Ramsay & Oxley, 1997. RosETTE PLANTS
This growth form gives páramo vegetation its distinctive character (see Fig. 18). Two kinds ofrosette
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Fig. 17. Adaptation to tire in Espeletia pycnophylla ssp. angelensis in Páramo del Angel (Carchi, Ecuador) at about 3500 m. A. Plants that have not been subjected to tire. Note the marcescent leaves along entire stem, and living leaves clustered at the distal end that protect the buds (center plan! is ca. 2.5 m tall). B. Plants that have been subjected to tire. Note how the lowermost lea ves ofthe plan! ha ve been bumed off and the stem is naked, but the apex has been protected by the distal living leaves and the plan! continues to grow (talles! plants are ca. 3.5 m tall). (Photos by J. L. Luteyn taken in 1997.)
plants, the stem rosette and the acaulescent rosette, have been described. The stem rosette, the columnar woolly type of rosette plant, is the most typical and well-known growth form of páramo. Members of the genus Espeletia (Asteraceae), the frailejón (literally, "big friar")-so called because of the grayish woolly coat of pubescence-are the classical example. These plants produce an erect, normally unbranched, thick-woody stem tightly encased by the dense bases of old leaves (Cuatrecasas, 1968). The erect stems may be as tall as 15 m in undisturbed páramo (Fig. 18B-C; Color Plate II, bottom left). Lateral inflorescences are produced from the single aerial meristem. Other examples of this growth form are the fern Blechnum schomburgkii (Biechnaceae) (Color Plate V, bottom left) and Plantago serie ea subsp. penymondii (Plantaginaceae). It has been shown that woolliness is a response to ultraviolet light and is associated with
thermoregulation, the retarding of evaporation, and general protection from UV light (Miller, 1994). Acaulescent rosette plants develop thick, perennial, taproots and a dense rosette of leaves at the ground leve!. The flowering stems may be very short, with the flowers hidden in and amongst the leafbases, or longer, thereby lifting the flowers above the ground surface. Sorne plants, such as Puya spp. (Bromeliaceae), produce giant, bulky inflorescences severa! meters tall (Fig. 18D). In all cases, the buds that form the flowers originate in the axils ofthe rosette leaves, which are sunken a few centimeters below the ground. Acaulcscent plants thereby protect their buds from fire and frost. Experiments ha ve shown that the mean temperature within the rosette is higher (by about 6°C) than in the surrounding air (Hedberg & Hedberg, 1979) and that these plants seem to buffer temperature variations to such an extent that they are able to avoid both positive and negative extremes (Goebel,
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Fig. 18. Rosette growth form. A. Coespeletia timotensis from the superpáramo ofPáramo de Piedras Blancas (Mérida, Venezuela) at ca. 4300 m. The plant at the center is about 2.3 m tal!. (Photo by F. L. Pérez taken in 1980.) B. Páramo de Santa Lucía, just south ofthe Laguna de La Cocha (Putumayo, Colombia) at ca. 3100 m. Subpáramo woodland with Espeletia sp. (For size comparison, note man alongside plant in lower center right.) (See Fig. 16A for details of associated plants.) (Photo Cuatrec. C1825 taken in 1941.) C. Espe!etia discoidea from Páramo de Guantiva (Boyacá, Colombia) at ca. 3250 m (H. García-Barriga is standing alongside the plant). This wet páramo al so containedEspelelia nemenkenii, Espeletia congestijlora, Espe/etia arbe/aezii, and Espeletiopsis muiska. Other associates included Ca/amagrostis planijolia, Blechnum sp., Rubus sp., Weinmanniafagaroides,
Pentacalia corymbosa, Val/ea stipularis, Be)aria aestuans, Gaultheria anastomosans, Pernettya pros/rata, Hypericum sp.,Excremis coarctata, and Paepalanthus crassicaulis. (Photo Cuatrec. C-1685 taken in 1940.) D. Puya goudotiana, an ac:aulescent rosette plant from Macizo de Bogotá region, in páramo near Guadalupe above Santafé de Bogotá (Cundinamarca, Colombia) at.ca. 3220 m. Al so in this community were Espeletia grandijlora, Espeletiopsis co¡ymbosa, Aragoa sp., Miconia pallida, Brachyotum sp., Monnina salicifolia, and Paepalanthus sp. (Photo Cuatrec. C-1390 taken in 1939.)
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1891; Hedberg, 1964). Examples are found in Hypochaeris (Fig. 13A) and Werneria (Asteraceae), Lysipomia (Campanulaceae-Lobelioideae), Paepalanthus (Eriocaulaceae) (Fig. 12A), Lupinus alopecuroides (Fabaceae) (Fig. 12C), Acaulimalva and Nototriche (Malvaceae), Rumex tolimensis (Polygonaceae), Ranunculús gusmanni (Ranunculaceae ), A caen a cylindristachya (Rosaceae), Valeriana plantaginea (Valerianaceae), and Viola (Violaceae). CusHION PLANTS
These plants form a flat, convex, or hemispherical cushion as the result ofthe regular outward branching of dense radially oriented buds (Fig. 19). Each branch has a small rosette ofleaves at the tip, and only the outer and upper leaves are green and living; the interior of the cushion consists of a peaty mass, the remains of dried leaves, accumulated humus, dust, soil, and rain water, all ofwhich protect the buds and stems from wind, desiccation, and predation, and pro vide a reservoir ofwater and nutrients (Sklenáf, 1998).1As in rosette plants, the mean temperature is higher within the cushion than at the cushion surface, thereby protecting the buds from cold temperatures. The plants are often very prickly and hard to the touch, yet firm enough that one is able to walk on top of certain species, such as in Azorella (Apiaceae), Plantago (Plantaginaceae) (Figs. 9C & 19), andDistichia (Juncaceae) (Fig. 15A). Many different species, in different families, form cushions of different sizes, from flat or only a few centimeters tall to cushions over one meter tall and severa] meters in diameter, e.g., Azore!!a pedunculata. In moist or humid sites, where cushions are more frequently found, Distichia muscoides and Plantago rígida are dominant species (Figs. 9C, 15A & 19; Color Plate II, top right), whereas in drier places Azore!!a pedunculata andA. aretioides are common. Other genera that produce cushions include Wemeria pygmaea andXenophyllum spp. (Asteraceae) (Color Plate VI, top left), Draba aretioides (Brassicaceae), Arenaría spp. (Caryophyllaceae), Oreobolus spp. (Cyperaceae),Disterigma empetrifolium and Pernettya pros trata (Ericaceae ), Paepalanthus lodiculoides (Eriocaulaceae), Geranium spp. (Geraniaceae), Aciachne pulvinata (Poaceae), Calandrinia acaulis (Portulacaceae), Valeriana rigida (Valerianaceae) (Color Plate V, bottom right), and Xyris subulata (Xyridaceae). For extensive discussions ofrnorphology and actaptation of cushion plants to hÍgh~eievation environments, see Hedberg, 1964, 1992; Heilbom, 1925; Rauh, 1939; and Sklenár, 1998. BUNCHGRASSES (oR TUSSOCK GRASSES)
This growth form is the most widespread in the páramo. In undisturbed areas, grasses may average 1-
23
1.5 m tall with a.coverage ofup to 100%. Members of the grass and sedge families frequently form tufts or dense bunches of stems (culms) with rigid, pointed, tubular or inrolled leaves (Figs. 8C,D, 9, 17; Color Plates I, bottom right and V, top right). These dense tufts in which the dead leaves are maintained and decay on the plant, along witlrthe culms, provide good insulation for the buds and young leaves from cold temperatures, high radiation, evaporation, and heat of fires to 500°C (Ramsay, 1992; Ramsay & Oxley, 1996). Much oftheir regeneration takes place through the production of vegetative buds near the ground. Here the tufts are very dense and living shoots are found along with dead culms and leaves. These tufts protect the vegetative buds. The most common species are Calamagrostis recta and C. effitsa (Poaceae) (Figs. 9B,C, 13D; Color Plate II, bottom left); other important genera and species are Carex and Uncinia (Cyperaceae), Cortaderia spp., Festuca dolichophylla, F. tolucensis (Fig. 9A), Stipa spp., and Lorenzochloa erectifolia (Poaceae). For additional discussion ofhow the bunchgrass growth form is an adaptation to fire in the páramo environment, see Hofstede, 1995c; Ramsay, 1992; and Ramsay & Oxley, 1997. MICROPHYLLOUS SHRUBS
These shrubs are characterized by dense foliage of small, xeromorphic leaves, sometimes with many of the following combinations of adaptations in the same species, al! acting as protection from ultraviolet light and/or the reduction oftranspiration (Hedberg, 1964; Larcher, 1975). Examples of genera and/or species with hard or sclerophyllous leaves include Gaultheria anastomosans and Gaylussacia buxifolia (Ericaceae) and Miconia summa (Melastomataceae); squamous or rolled leaves Baccharis revoluta and Diplostephium revolutum (Asteraceae) and Miconia salicifolia (Melastomataceae); imbrica te lea ves Loricaria (Asteraceae) (Color Plate V, top left) andAragoa cupressina (Scro. phulariaceae ); aciculate or spine-tipped lea ves Chuquiraga (Asteraceae) (Fig. 11D), Hypericum laricifolium (Clusiaceae), and Valeriana microphylla (Valerianaceae ); and densely tomentose-pubescent leaves Diplostephium eriophorum and Pentacalia guicanensis (Asteraceae) and Gaultheria lanigera (Ericaceae). The so-ca1led bouquet plants ("plantas en ramilletes de florones") are a subset of microphyllous shrubs with a special habit. These plants feature an increase in the size of an individual flower in relation to the total appearance of the plant (e.g., Bidens hu milis), or the dense aggregation of many small flowers into a bouquet of dense flowering stalks (Vareschi,
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Fig. 19. Pรกramo community showing cushion plant growth form. Climbing Nevado del Ruiz abo ve Termales (Caldas, Colombia) looking to the East at ca. 3900 m. In the foreground are f01mations ofthe bunchgrasses Calamagrostis ejjitsa and C. recta, and cushion plants of Plantago rigida (From J. Cuatrecasas, "Aspectos de la vegetaciรณn natural de Colombia," Revista Acad. Colombiana Ci. Exact. l 0(40). Reproduced with permission. Photo Cuatrec. I-235 taken in 1940.)
1970). This is considered an adaptation that makes the flowers more attractive to pollinators by forming a large and noticeable splash of color that attracts insects from a distance, as in species of Draba (Brassicaceae), Gentianella (Gentianaceae), and Chaetolepis (Melastomataceae) (Color Plate III, top left). PROSTRATE DWARF SHRUBS
These are small, woody plants that rarely produce shoots over 0.75 m tall. The speciaร feature separating them from microphyllous shrubs is that they ha ve a larger part of their branch system protected below or on the soil surface. They are often pros trate, growing laterally along the ground. Sometimes entire
branching systems occur underground and only the current year's growth is seen above ground. This growth form often has its regenerative buds below ground, where they are protected from fire and frost. Examples are found in Bidens and Senecio (s.l.) (Asteraceae), Lupinus and Astragalus (Fabaceae) (Color Plate VI, bottom left), Pernettya andDisterigma (Ericaceae), andArcytophyllum (Rubiaceae). GEOPHYTES
These are herbs that survive the unfavorable periods of the year (including times of fire) by means of subterranean organs, such as succulent roots, rhizomes, stolons, tubercules, or bulbs (L<egaard, 1992; Raunkiaer,
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1934; Vareschi, 1970). Examples of geophytes include Orthrosanthus chimboracensis (Iridaceae) (Color Plate VI, top right), the genus Stenomesson (Amaryllidaceae), Altensteinia and Gomphychis (Orchictaceae), and the fem Ophioglossum crotalophoroides (Ophioglossaceae).
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Table I Numbers offamilies, generá, and species ofmajor plant groups in the páramo Taxanomic group Non-vascular plants
V AGRANT PLANTS These plants grow free, unattached to the substrate and are found in many biomes throughout the world (Pérez, 1994a, 1997b ). They are found in the superpáramo zone, where frost-heaving is a common phenomenon, and only in cryptogamic plants such as the fruticose lichen Thamnolia vermicularis (family uncertain) and the acrocarpous moss Grimmia longirostris (Grimmiaceae). These plants have been variously referred to as erra tic, vagant, vagrant, solifluction floaters, errant cryptogams, and sometimes globular mosses or moss balls when their growth shape becomes more spherical (see Pérez, 1997b, for more details). Cryptogams as a whole (i.e., bryophytes and lichens) have been considered a true growth fonn by Cuatrecasas (1968), Griffin (1979), and Ramsay (1992), oras composed of severa] growth forms by Ramsay and Oxley (1997).
Flora On a geological timescale the páramo flora is young, the so-called protopáramo vegetation of Van der Hammen and Cleef (1986) having evolved during the Late Pliocene or Early Pleistocene, sorne 2~ Ma. Páramo environments suitable for plant colonization on an extensive scale, however, have been available only since the Quatemary (Simpson, 1975; Van der Hammen, 1974; Van der Hammen & Cleef 1986). The youth ofthe páramo flora is al so evidenced by the presence of relatively few endemic or near endemic genera (23 genera, ca. 5% based on this work) and the absence of any endemic families in the vascular flora. Although the páramo ecosystem occupies no more than 2% ofthe land area ofthe countries in which it is found, the flora is extremely diverse. In fact, the páramo flora is the richest high-mountain flora ofthe world (Smith & Cleef, 1988). The páramo flora has evolved in various ways: by adaptation of lower-elevation plants (i.e., tropical elements) to highelevation environments, by immigration (i.e., dispersa!) of cool-adapted plants from north and south temperate regions, and by speciation through isolation from within (i.e., autoc)1thonous element). For discussion of these and other ideas, see Chapman, 1917; Chardon, 1938; Simpson, 1975; Simpson & Todzia, 1990; Van der Hammen, 1972a, 1972b; and Vuilleumier, 1970.
Lichens' Mosses Hepatics Vascular plants Fems/Fem Allies
Families
Genera
Species
130 45 51 34 124 22
365 114 163 500 52
101 16 85 254
447 101 346 865
1298 465 544 291 3399 352 2 3045 634 2411 4697
88
Gymnosperms Angiosperms Monocots Dicots Total
'Lichenicolous fungi are not included in this table.
GENERAL FLoRrsnc DrvERSITY
For the nonvascular plants, the lists herein presented include 114 genera in 45 families of lichens (excluding lichenicolous fungi), 163 genera in 51 families of mosses, and 88 genera in 34 families of hepatics, for a total of 365 genera and 1298 species (see Table I). For the vascular plants, the lists in elude 52 genera in 22 families of fems and fern allies 1 genus in 1 family of gymnosperms, 101 genera in '16 families of monocotyledons, and 346 genera in 85 families of dicotyledons, for a total of 500 genera and 3399 species (see Table I). Tables II-VI show the largest families and genera of páramo lichens, mosses, hepatics, ferns and fern allies, and flowering plants, respectively. Table VII lists the genera endemic to páramo and notes those that are monotypic. Luteyn (1992) provided a preliminary estima te of specific endemism of páramo vascular plants as high as 60%. However, it is now realized that species numbers, limits, and distribution patterns are too poorly known and that a realistic approximation of species endemism in the páramo is not yet possible. As a result, detailed estimates of overall specific endemism and the geographical origins and relationships ofthe páramo flora have not been calculated from the data herein presented. Nonvascular plants.-In this study, 1298 species of non vascular plants have been found throughout the geographical and elevational range ofpáramo as here defined (Table I), of which 36% are lichens (lichenicolous fungi are not included) 42% mosses and 22% hepatics. ' '
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Table U Ten largest families and genera ofpáramo lichenized fungi• (prepared by H. Sipman) Fami1y (no. generalspecies) Parme1iaceae (25/159) C1adoniaceae (3/45) Physciaceae (8/27) Collemataceae (2/26) Lobariaceae (3/24) Stereocau1aceae (1/19) Peltigeraceae (3/13) Rama1inaceae (1/12) Lecanoraceae (4/11) Pannariaceae (4/1 O)
Genus (fami1y) (no. species)
Hypotrachyna (Parme1iaceae) (50) Cladonia (C1adoniaceae) (38) Leptogium (Collemataceae) (25) Oropogon (Parmeliaceae) (21) Stereocau/on (Stereocau1aceae) (19) Xanthoparme/ia (Parme1iaceae) (18) Heterodermia (Physciaceae) ( 17) Sticta (Lobariaceae) (13) Parmotrema (Parmeliaceae) (13) , Ramalina (Rama1inaceae) (12)
'Lichenico1ous fungi are not included in this tab1e.
Table 111 Ten largest families and genera ofpáramo mosses (prepared by S. P. Churchill and D. Griffin III) Fami1y (no. generalspecies)
Genus (fami1y) (no. species)
Dicranaceae ( 17/67) Bryaceae ( 10/65) Pottiaceae (19/63) Bartramiaceae (7 /40) Orthotrichaceae (3/36) Sphagnaceae (1/27) , Amblystegiaceae (9/19) Brachytheciaceae (7 /18) Grimmiaceae (4/ 17) Polytrichaceae (6/16)
Campylopus (Dicranaceae) (37) Sphagnum (Sphagnaceae) (27) Zygodon (Orthotrichaceae) (21) Bryum (Bryaceae) (18) Leptodontium (Pottiaceae) ( 16) Orthotrichum (Orthotrichaceae) (14) Breutelia (Bartramiaceae) ( 13) Daltonia (Daltoniaceae) (13) Macromitrium (Macromitriaceae) (13) Schizymenium (Bryaceae) (11)
Within the lichens, Table II shows the 10 largest families and genera of páramo lichenized fungi. At the fami1y leve!, Parmeliaceae (25 genera and 159 species) are by far the most diverse, with more than three times as many genera and species as the next closest families Physciaceae (8 gen.) and Cladoniaceae (45 spp.), respectively. Four of the 10 most speciose páramo genera are al so ParmeliaceaeHypotrachyna (50 spp.), Oropogon (21 spp.),Xanthoparmelia (18 spp.), and Parmotrema (13 spp.)-accounting for just over 100 species. The second most speciose family is Cladoniaceae, with Cladonia having 38 species. Additional general comments about lichens are given by Ahti (1992) and Sipman (1992, and below in the introduction to his checklist). Table III shows the 1Olargest faínilies and genera ofpáramo mosses. At the family leve!, Dicranaceae (17 gen., 67 spp.), Bryaceae (10 gen., 65 spp.), and Potti, aceae (19 gen., 63 spp.) are the most diverse. The most speciose genera are Campylopus (Dicranaceae, 37 spp.),
Sphagnum (Sphagnaceae, 27 spp.), and Zygodon (Orthotrichaceae, 21 spp.). Additional general comments about mosses are given below by Churchill and Griffin in the introduction to their checklist. Table IV shows the 1O largest families and genera of páramo hepatics. Lejeuneaceae (16 gen., 38 spp.) and Jungermanniaceae (11 gen., 31 spp.) are the most diverse families in terms ofboth genera and species. The most speciose genera are Riccardia (Aneuraceae, 20 spp.), Metzgeria (Metzgeriaceae, 20 spp.), Plagiochila (Plagiochilaceae, 18 spp.), Frullania (Jubulaceae, 13 spp.), and Bazzania (Lepidoziaceae, 13 spp.). Additional general comments about liverworts are given below by Gradstein in the introduction to his checklist. Vascular plants.-In this study, 3399 species ofvascular plants have been found from throughout the geographical and elevational range of páramo (Table I), of which 10.4% are fems and fem allics, 0.06% gymnosperms, and 89.6% angiosperms. Of the angiosperms (flowering plants), 21% are monocots and 79% are dicots.
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Table IV Ten largest families and genera of páramo hepatics (prepared by S. R. Gradstein) Fami1y (no. genera/species)
Genus (family) (no. species)
Lejeuneaceae (16/38) Jungermanniaceae (11/31) Lepidoziaceae (6/29) Aneuraceae (2/21) Metzgeriaceae (1/20) Plagiochilaceae (2/19) Geocalycaceae (7/18) Gymnomitriaceae (5/14) Jubulaceae (1/13) Ba1antiopsaceae (2/7)
Riccardia (Aneuraceae) (20) Metzgeria (Metzgeriaceae) (20) Plagiochila (P1agiochi1aceae) (18) Frullania (Jubu1aceae) (13) Bazzania (Lepidoziaceae) (13) Anastrophyllum (Jungermanniaceae) (8) Lepidozia (Lepidoziaceae) (8) Leptoscyphus (Geoca1ycaceae) (7) Isotachis (Balantiopsidaceae) (6) Cephaloziella (Cephaloziellaceae) (6) Marsupe/la (Gymnomitriaceae) (6) Radula (Radulaceae) (6)
Table V Ten largest families and genera ofpáramo fems and fem allies Family (no. genera/species)
Genus (family) (no. species)
Dryopteridaceae (5/77) Lycopodiaceae (3/69) Polypodiaceae (14/60) Pteridaceae (7 /43) Hymenophyllaceae (1/20) Isoetaceae (1/18) Thelypteridaceae (1/13) Aspleniaceae (2/11) Blechnaceae (1/9) Cyatheaceae ( 1/6)
Elaphoglossum (Dryopteridaceae) (65) Huperzia s.str. (Lycopodiaceae) (60) (Lycopodium s.l. = 69) Hymenophyllum (Hymenophyllaceae) (20) Isoetes (Isoetaceae) (18) Jamesonia (Pteridaceae) ( 18) Eriosorus (Pteridaceae) (14) Thelypteris (The1ypteridaceae) (13) Polypodium (Polypodiaceae) (12) Melpomene (Polypodiaceae) (11) Asplenium (Asp1eniaceae) ( 1O)
Table V shows the largest families and genera within the páramo ferns and fem allies. Dryopteridaceae, Lycopodiaceae, Polypodiaceae, and Pteridaceae are the largest and most di verse families at the generic and specific levels. At the generic leve!, Polypodiaceae (14 gen.) are the most diverse, with twice as many genera as the next closest family, Pteridaceae (7 gen.). At the specific leve!, Dryopteridaceae are the largest dueto the numerous species of Elaphoglossum (65 spp.); this genus is also the largest ofthe páramo pteridophytes. Huperzia (Lycopodiaceae), with 60 species, is the second largest genus (or the largest, with 69, if considered as Lycopodium in the broad sense), followed by Hymenophyllum (20 spp.), Isoetes (18 spp.), and Jamesonia (18 spp.). Table VI shows the composition of the páramo flowering plants in terms ofthe 15largest families and genera. In this study, the Asteraceae are the largest family by far in both numbers of genera and species. The data from this study show that Asteraceae are two
and a half times larger than the Poaceae in numbers of genera (101 gen. vs. 41 gen.) and nearly four times larger in numbers of species (858 spp. vs. 227 spp.). Four of the five most speciose páramo genera are Asteraceae-Pentacalia (89 spp.), Senecio s.str. (69 spp.), Diplostephium (70 spp.), andEspeletia s.str. (61 spp.)-accounting for nearly 300 species. Asteraceae also have the highest number of endemic genera of any páramo vascular plant family ( 16 genera, or 70% ofthe endemic páramo genera). The largest genera of Poaceae are Festuca (38 spp.), Calamagrostis (36 spp.), Agrostis (24 spp.), and Poa (20 spp.). The Orchidaceae are surprisingly diverse in the páramo, with 25 genera and 152 species herein recorded, al" though (as discussed below) their tabulation has pro ven difficult and their numbers may be questioned. It is interesting to note that in virtually all páramo studies that have been published, the Asteraceae are always the largest family in numbers of species and genera, followed closely by Poaceae. Other families
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Tablé VI Fifteen largest families and genera ofpáramo flowering plants Fami1y (no. genera!species) Asteraceae (101/858) Poaceae (41/22 7) Orchidaceae (251152) Scrophulariaceae (14/144) Melastomataceae (9/1 07) Gentianaceae (4/93) Ericaceae (16/79) Bromeliaceae (6/78) Rosaceae (1 0/77) Fabaceae" (9/76) Brassicaceae (13/71) Cyperaceae (8/70) Apiaceae (15/61) Solanaceae (8/58) Clusiaceae (2/56)
Genus (fami1y) (no. species) Pentacalia (Asteraceae) (89) Senecio s.str. (Asteraceae) (69) (Senecio s.l. = 172) Diplostephium (Asteraceae) (70) Ca/ceo/aria (Scrophu1ariaceae) (65) Espe/etia s.str. (Asteraceae) (61) (Espe/etia s.l. = 123) Lupinus (Fabaceae) (56)' Miconia (Melastomataceae) (54) Hypericum (C1usiaceae) (54) Va/er¡'ana (Valerianaceae) (54) Gen.tianella (Gentianaceae) (48) Puj;a (Bromeliaceae) (48) Gynoxys (Asteraceae) (46) Baccliaris (Asteraceae) (45) . Draba (Brassicaceae) (45) Geranium (Geraniaceae) (43) Soianum (Solanaceae) (43)
"Rupert Bameby, who is curren ti y studying Lupinus, feels there are only about 15 species in the genus. Ifthis is true the overall family number drops toca. 35 species.
Table VII Vascular plant genera endemic (or nearly so) to the páramo. Those marked with an asterisk (*)are monotypic Vascular plants Apiaceae Cotopaxia Perissicoe/um Asteraceae Aphanactus Ascidiogyne
Blakiella' C!uysactinium Coespe!etia Espeletia Espe!etiopsis Floscaldasia' F/osmutisia'
Freya Hinterhubera Ja/cophila La estadía Paramiflos Raouliopsis Westoniel/a
that rank consistently high in overall importance in the páramo ecosystem include (in alphabetical order) Apiaceae, Brassicaceae, Bromeliaceae, Cyperaceae, Ericaceae, Gentianaceae, Melastomataceae, Orchidaceae, Rosaceae, and Scrophulariaceae. There is also a trend in the upper páramo (i.e., superpáramo at ca. 4000+ m) for certain families, often with a more northtemperate element, to become increasingly importan!. For example, the Apiaceae, Brassicaceae, Caryophyllaceae, Fabaceae, Gentianaceae, Malvaceae, Poaceae, and Valerianaceae become more conspicuous, while the Bromeliaceae, Melastomataceae, Orchidaceae, Rubiaceae, and Solanaceae decline (see also Jorgensen & Ulloa U., 1994: tab. 9). GENERAL PHYTOGEOGRAPHY ANO ÜRIG!NS
Frequently, the overall floristic and phytogeo-
Campanulaceae Lysipomia Melastomataceae Castratella Scrophulariaceae Aragoa
Ferns Pteridaceae Jamesonia Nephopteris'
graphical relationships ofthe páramo flora have been discussed and compared with the Mexican and Guatemalan alpine floras (Beaman, 1965; González, 1986; Islebe & Cleef, 1995; Rzedowski, 1978), the tepuis ofthe Guayana High1ands (Cleef et al., 1993; Riina, 1996), the lowland savannas of South America in general (Cleef et al., 1993), the puna flora (Baumann, 1988; Quintanilla P., 1983b), the subantarctic flora (Cleef, 1978, 1980a), and the world's other high-elevation tropical floras such as in A frica and Malesia (Smith & Cleef, 1988). In common with al! other highelevation tropical floras, the páramo flora is predominantly of temperate zone origin at the generic leve]; but in contrast, the páramo flora is the richest overall and has the largest actual number of genera and endemic elements (Smith & Cleef, 1988). Future research efforts are needed to see ifthe previous com-
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Table VIII Percentage ofvascular plant genera in each phytogeographical element in the neotropical páramos
Geo¡¡;raEhical element Páramo Qther neotropical Wide tropical Holarctic Austral-Antarctic Wide temperate CosmoEolitan
CR' 4 25 7 15 14 24 11
Percentage ofvascular ¡;¡lant genera VEh C0-2d EC-1' CO-l' 7 8 4 7 24 34 32 30 3 10 28 10 13 ll 12 10 7 9 5 10 39 26 20 7 7 8 10 9
EC-2' 9 21 3 14 10 26 17
'Cordillera de Talamanca, Costa Rica (Cleef & Chaverri P., 1992). hBased on data from the superpáramo ofPáramo de Piedras Blancas, Venezuela (Ricardi S. et al., 1997). 'Based on ca. 600 relevés from all parts of the páramo throughout the Cordillera Oriental, Colombia (Cleef, 1979a). dBased on the 130 most important species of the Colombian páramo flora as a whole (Sturrn & Rangel Ch., 1985). 'Páramo de Guamaní, Ecuador (León Y., 1993). 'Based on 192 quadrats in zonal vegetation from 12 scattered páramos in Ecuador (Ramsay, 1992).
parisons, based mostly on a restricted data-set from the páramos ofthe Colombian Cordillera Oriental, are representative for the páramo flora as a whole. In an effort to understand the origins ofthe páramo flora, recent phytogeographical studies have followed Cleef(1979a) in assigning each genus to one ofseven geographical floristic elements: páramo, other neotropical, widespread tropical, Holarctic, AustralAntarctic, widespread temperate, and cosmopolitan (see Table VIII). In the Colombian Cordillera Oriental, the most comprehensively studied páramo region to date, Cleef (1979a, 1980c) and Van der Hammen and Cleef (1986) found that the páramo genera of vascular plants are about 50% tropical in origin and 50% temperate. Within the tropical element, strictly neo tropical genera are the most strongly represented. Within the temperate element, the widespread genera are most strongly represented (about 20% ofthe genera), while the Austral-Antarctic and Holarctic elements are represented each by about 10%. See al so Sturm and Rangel Ch. (1985), who studied the phytogeography ofthe 130 most important species ofthe Colombian páramo flora as a whole. In the páramo flora ofthe Cordillera de Talamanca (Costa Rica), Cleef and Chaverri P. (1992) found that, of the páramo genera of vascular plants, about 36% are tropical in origin and 64% temperate. Within the tropical and temperate elements ofCosta Rica, strictly neotropical genera and widespread genera, respectively, are the most strongly represented, as they are in the Colombian Cordillera Oriental. Although the Costa Rican páramo flora shares about 95% of its
vascular genera with the Andes, the larger proportion ofthe temperate component in Costa Rica, with genera such as Garrya (Garryaceae), Arctostaphylos (Ericaceae), Mahonia (Berberidaceae), and Romanschulzia (Brassicaceae), was attributed to the more northern geographical position ofthe country. [Note: The genera Helianthemum (Cistaceae) and Smilacina (Liliaceae) have also been reported from the páramos of Costa Rica by Cleef and Chaverri P. (1992), but I have not seen any herbarium specimens unequivocally from páramo and, therefore, ha ve not included them in this book.] Ricardi S. et al. (1997b) studiedthe superpáramo (above 4000 m) in Venezuela and compared it with other high-elevation floras along the entire length of the Andes from Costa Rica to Patagonia. They found that 34% ofthe genera ofvascular plants are tropical in origin and 66% temperate, with the greatest similarity to that ofColombia and next to Ecuador. In northern Ecuador, León Yánez (1993) found that, ofthe genera ofvascular plants, 45% are oftropical origin and 55% of temperate. In the southern páramos of Ecuador, Ramsay (1992) found fewer families and genera than in Colombia, with about 33% ofthe genera ofvascular plants oftropical origin and 67% temperate. He attributed this to the lower humid~ ity and more extreme cold temperatures ofEcuadorean páramos. Ramsay (1992) also compared the Ecuadorean páramo flora to the puna ofPem and the mountains ofEast Africa and New Guinea (see also Balslev, 1988; Mena V., 1984; andMena V. &Balslev, 1986). There are, unfortunately, no similar phytogeographi-
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cal studies from the páramos (jalea) ofnorthem Petu. phibians such as the salamander (Bolitoglossa spp.), Monasterio (1980b) looked at the entire páramo frog and toad (Eleutherodactylus spp., Hyla spp., region and found the following trend: the jalea and Ate/opus spp.), and reptiles such as the lizard and Ecuadorean páramos consist mainly oftussock grasses chameleon (Stenocercus spp., Phenacosaurus spp., with genera from extra-tropical regions, the Venezu- Proctoporus spp.). Invertebrates are less conspi'cuous and found elan páramos are dominated by rosettes, and the páramos of Colombia show equal importance of mostly in the subpáramo. These include lepidopterans (Descimon, 1986) and other insects such as grassgrasses and rosettes. For additional studies that include phytogeographi- hoppers, cockroaches, beetles, and flies; molluscs cal analyses, see Ahti, 1992 (lichen family Cladoni- such as a few snails and slugs; and earthworms. aceae); Becker, 1988; Cabrera, 1957; Cleef, 1981b; Arthropods and other microfauna (e.g., mites and Duque N., 1987; Jorgensen & Ulloa U., 1994; springtails) are rarely seen but are abundant (Sturm, Keating, 1995; Lozano C. & Rangel Ch., 1989; Rangel 1978, 1983, 1990, 1994a; van VeJez, 1992). Ch., 1991b, 1995a; Rangel Ch. & Garzón, 1995, 1997; . A number ofthe larger páramo mammals, as well Sipman, 1992 (Colombian lichens); Tirado M. & · as the Andean candor, ha ve been extensively hunted Ricardi S., 1997); Vargas Ulate & Sánchez G., unpubl. by man and their numbers are now very low. Hum. mingbirds, bees, and flies seem to be important polliFauna nators in the páramo (see, e.g., Berry & Calvo, 1994; Brand Prada, 1994b). Birds, rabbits, and guinea pigs Páramo grass- and shrublands support a number (as well as wind, water, and gravity) are important of native animal species. It is not the purpose of this agents of dispersa! (Frantzen & Bouman, 1989; Graf book to discuss the general fauna ofthe páramos, but Bock, 1984; Simpson & Todzia, 1990). sorne of the larger, more common, and conspicuous Additional references about the páramo fauna include animals are mentioned here. Aagaard, 1982; Adams, 1973; Amat García, 1987, Mammals in the páramo include the puma (Felis 1991a, 1991b; Arias Lemas, 1989; Aristide U., 1969; concolor), spectacled bear or "oso de anteojos" Bamett & Gordon, 1985; Barrientos & Monge-Nájera, (Tremarctos ornatus), white-tailed deer (Odocoileus 1995; Berna] C., 1985; Berna! C. & Figueroa, 1980; virginianus), mountain tapir or "danta". (Tapirus Brand Prada, 1994a; Cadena G. & Malagón, 1994; pinchaque), rabbit (Sylvilagus brasiliensis), guinea pig Chapman, 1917, 1926; Davis et al., 1997; Del Llano, (Cavia porcellus ), Andean fox (Duscicyon culpaes ), 1990; Duellman, 1979a, 1988; Fjeldsil. & Krabbe, 1990; mountain coatí (Nasuella olivaceae), long-tailed weaGaray, 1981; E. Gaviria, 1991; S. Gaviria, 1989; sel (Mustela ji-enata ), shrew ( C!yptotis spp. ), rat oposGrabandt, 1983; Hoffstetter, 1986; Hoyos, 1991; sum (Caenolestes spp.), and various small rodents. INDERENA, 1984; Janzen et al., 1976; Lynch, 1986; Rabbit feces are extremely common in the grass Maní, 1962; Mora O. & Sturm, 1994a; Phelps & Phelps, páramo, especially in open areas between tussocks. 1958, 1963; Rangel Ch. & Berna!, 1980; Reig, 1986; Stotz et al. (1996) list 69 species ofbirds as "total Righi, 1995; Rivera, 1979; Sturm, 1984a; Sturm & users" ofthe páramo habitat, with 41 species making Rangel Ch., 1978; Vuilleumier, 1980; Vuilleumier & it their primary habitat and 16 as indicator species. Ewert, 1978; Wolf & Gil!, 1986; and Yerena, 1994. Sorne ofthe better-known páramo birds include vultures su eh as the Andean candor ( Vultur gryphus) and Human Influence the turkey buzzard ( Cathartes aura), eagle, hawk, and falcon (Phalcoboenus carunculatus, Buteo spp., There have been settlements in the highlands of Geranoaetus spp., Falca sparverius), owl (Buba virgi- the Andes for perhaps 15,000 years or more (Eckholm, nianus andAsio spp.), hummingbird or "colibrí" (fam1975; Little, 1981). Man's influence there has been ily Trochilidae: e.g., Oxypogon guerinii and Patagona profound, with the result that nearly 90-95% of the gigas), duck (A nas spp.), and the rufous-fronted para- forests of the northern Andes have been cleared keet (Bolborhynchus ferrugineifi·ons). For a discus- (Henderson et al., 1991). Exploitation ofthe high-elsion ofthe origins ofthe high Andean bird fauna, see evation puna ecosystem of the central and southern Vuilleumier, 1986. Andes of Peru, Bolivia, and northern Argentina ocNative páramo fish species are few, but rainbow trout curred in the pre-Hispanic period and has been relaor "trucha" (Oncorhynchus mykiss) have been introduced tively well documented (see, e.g., Baker & Little, to many lakes and streams for food and sport. 1976; Brush, 1976, 1982; Ellenberg, 1979). Intensive The herpetofauna has been relatively well studied land use in the páramos ofthe northern Andes has been (see especially Duellman, 1979b) and includes am- a more recent phenomenon and needs further study
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(Hess, 1990). Did similar changes ofhabitat destruction and alteration take place in the essentially unpopulated high-elevation páramo areas as it did in the puna or in the lower montan e regions of the Andes? One of the most frequently discussed questions about the historical development of highland regions ofthe northem Andes is whether the páramo that we see today, specifically the grass páramo, is a natural ecosystem or one that man created in order to provide more pasture and agriculturalland. Ellenberg (1979) championed the theory that the climax vegetation ofthe tropical Andes was forest and that man has been responsible, largely by means of fire, for nearly all the large open areas (Becker, 1988; Fjeldsa, 1992;-La::gaard, 1986, 1992). Other authors feel that both Polylepis forest and sorne form of páramo grassland have always existed as independent formations, expanding and contracting throughout time in response to climatic changes (Cleef, pers. comm.; Lauer, 1981; Troll, 1959; Van der Hammen & Cleef, 1986; Walter & Medina, 1969a, 1969b). Charcoal fragments in core sediment samples revea! that fires have occurred in-páramo since Holocene times (Hom, 1989c; Hom & Sanford, 1992; Salomons, 1986). Those fires may have been ignited by natural Sources (i.e., lightning or volcanic activity) or by man (deliberate or accidental), but usually their exact origins are unknown. Verweij (1995) has stated that fires caused by natural sources occur less than once in 1000 years. Published records oflightning-set páramo fires have not been found, although Horn (1989c) and Young (pers. comm.) ha ve mentioned lightning strikes in Chirripó páramo (Costa Rica) and the Río Abiseo area (northern Peru), respective! y. In studies of the Costa Rican páramos, Hom (1989c, 1993, in press; Hom & Sanford, 1992) has correlated modem and prehistoric fires together with palien analyses of sediment cores. She has concluded that in the Chirripó highlands páramo communities are the "natural" vegetation and that modem poli en assemblages of individual taxa have not differed greatly from those developed since deglaciation sorne 10,000 years ago. She has also found that fires have been a part ofChirripó páramos throughout the Holocene but have not "carved páramo from forest." Without doubt man has had major impact on the origin and subsequent spread of grasslands throughout the Andes, and perhaps he has been the single most important reason why grass páramo exists today, where shrub/tree woodlands of Polylepis, Buddleja, and Gynoxys may have once dominated. It is unlikely, however, that we wili ever be able to say with confidence what percentage oftoday's páramo has anthrpogenic origins. Whateverthe outcome ofthis discussion,
31
the facts remain that grass páramo currently exists, covers large expanses of the high-elevation Andes, and has great ecological and economic importance. The following discussion summarizes what we do know about the history of man's presence in the páramos of the high Andes and the effect he has had on them. Although many ofthe references and quotes given below are specific to one country, I am certain that similar events have occurred in al! of the north Andean countries including Venezuela, Colombia, Ecuador, and probably, by extension, Costa Rica and Panama. The situation in southern Ecuador and northem Peru is less clear, however, and may be quite different due to its overall drier climate and proximity to the larger pre-Columbian civilizations that occupied high elevations to a much greater extent in the central Andes than in the northern Andes. For example, the intense grazing pressure may have occurred earlier in Peru because of the presence of native camelids and their herding. THE PRE-COLUMBIAN PERIOD
It is assumed that the Venezuelan páramos have been exploited for more than 1400 years (López del Pozo, 1992). They ha ve been occupied for at least 500 years, although there were no permanent settlements untll colonial times (Wagner, 1978, 1979, 1988). The páramos were used, however, as corridors, in rituals, oras hunting areas (Clarac, 1981, 1985; Rojas, 1985). According to Salgado-Labouriau (1976) and Monasterio (1980d), pre-Columbian agriculture was present in Venezuela up to 3000 m, the lower edge of the páramo, but it was concentrated below 2000 m and the cultures that developed in or near páramo were dedicated to agriculture and hunting/gathering (Molinillo & Monasterio, 1997). In Colombia, the pollen record shows that human activity in the páramos around Bogotá began about 800 years ago (Van der Hammen, 1968). The Colombian páramos were mostly uninhabited, however, although they too were used for the cultivation of sorne tuber crops and as a crop-storage area (Langebaeck, 1988; López del Pozo, 1992). The highestmountains and páramos were mostly considered sacred and had mystical qualities attributed to them; their main use was for religious purposes and to bury the prominent dead (G. Correal U., cited in Cleef, 1981b; Reichel-Dolmatoff, 1982; Zambrano, 1993). THE COLONIAL ANO lNDEPENDENCE PERIODS
Europeans settled the New World in the 16th century. They introduced exotic animals that were totaliy new to the northem Andes, viz., cows, sheep, goats,
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horses, and donkeys, which added to the herds of native THE MüDERN PERIOD During the 20th century, man's impact on páramo camelids (llama, alpaca, and vicuña) in Peru and further south. [Llamas and alpacas have only recently been in- has been a story driven by demographics, i.e., the troduced into the Ecuadorean páramo (Hervas Ordoñez, numbers, density, and distribution ofpopulations (see 1994).] Exotic plants were also introduced by Europe- Fig. 20). As population densities in the interandean ans for cultivation at the cooler high elevations, espe- valleys increased, people began to colonize new arcially the grains wheat and barley, but also vegetable eas in order to satisfy their needs (Bernsen, 1991; crops such as broad or faba beans, lentils, peas, carrots, Brush, 1976, 1982; Ellenberg, 1979; Hess, 1990; radishes, onions, and garlic. These new plants added to Hofstede, 1995c; López del Pozo, 1992; Molinillo & Monasterio, 1997; Monasterio, 1980d; Parsons, 1982; the native grain quinoa and tuber crops such as potato. Early colonists (including pre-Columbian man) Pérez, 1992a). The annual population growth rates, founded their villages and towns primarily in the previously over 3% in Venezuela, Colombia, Ecuacooler interandean valleys along rivers, where their dor, and Peru (Little, 1981) and currently 2.6-2.9% crops were grown in the fertile floodplains (com) and (World Population Data Sheet 1997 from the Populaalong the adj acent upland slopes (potatoes and grains). tion Reference Bureau), are still the highest in South Cattle reached the Andes of Venezuela and Colom- America. The Colombian population, for example, bia in the late 16th century but probably were not in- increased from 3 million to 12 million between 1900 troduced into the páramos until the early 18th, and and 1950, rose to 26 million in 1975 and then to 33 then only at elevations to ca. 3000 m (Vila, 1962; million in 1990, and is expected to reach 49 million Wagner, 1967). The cattle foraged in the páramo in 2025; the Venezuelan population increased from 5 grasslands above the forest, severa! hours' or days' million to 20 million between 1950 and 1990, roseto walk from their homes. As herds of cattle grew in 22 million in 1995, and is expected to reach 35 milnumbers, fires were periodically set to bum off the lion in 2025; while the Ecuadorean population indead material from the páramo and provide fresh new creased from 3 million to 11 million between 1950 grass for the cattle. With time the páramo became and 1990, rose to 12 million in 1995, and is expected overgrazed and eJ5.cessively bumed. W ood was also to reach 18 million in 2025 (W orld Resources, 1996cut from nearby forests, mostly for fue! and as build- 1997). ing material (including road-building). Gradual! y, the Because ofland and agrarian reforrn policies durneed arose to cut and burn the forest immediately ing the 1960s and 1970s, most parts of Colombia, below the páramo to provide additional new grass- Venezuela, Ecuador, and northern Peru have experilands for pasturing and arable land for crops. Sedi- enced two recent tendencies: the upward movement ment cores from Laguna Victoria, Venezuela (3250 m), of agriculture into the páramo belt between 3000 m show that at about this same time large amounts ofpol- and 4000 m, and the intensification of animal produclen of Rumex acetosella, an herb introduced with wheat tion in the lower páramo belts (Baruch, 1979; Becker, and potatoes, as well as a drastic drop in tree pollen, 1988; Bernsen, 1991; Ferwerda, 1987; Hess, 1990; suggesting a massive deforestation of the upper López-Zent, 1993; Monasterio, 1980d; Pérez, 1992b; Andean forest (Salgado-Labouriau & Schubert, 1977). Verweij, 1995; Verweij & Beekman, 1995). In VenThe numbers of cattle continued to rise steadily ezuela, Monasterio ( 1980d) reported that sin ce the during the 18th century. In the 19th century, during 1960s, the highest limit of cultivation (in Páramo the wars ofindependence and civil strife in Venezu- Piedras Blancas, ca. 4200 m) has increased dueto the ela (and possibly throughout Colombia, Ecuador, and introduction offrost-resistant varieties ofpotato, such Peru), the numbers of cattle oscillated, sometimes as the "papa negra." decreasing by almost 95%, although they again The last 20-25 years have also seen, however, a reached the pre-independence levels by ca. 184 7 regional growth in urbanization, with farrners aban(Brito, 1972; Pérez, 1992a, in press; Rouse, 1977; doning the countryside to cities such as Mérida, Vila, 1962, 1978). Bogotá, Nariño, Quito, Cajamarca, etc. This has reThus it would seem that in the early 18th century sulted in greater demand and increased pressure on the páramos ofthe northern Andes were still not utilized fewer farrners throughout the Andes to cultivate new to the extent ofthe altiplano orpuna to the south (Brush, land to produce more food. As natural soil fertility 1982). However, during the late Colonial period and decreased in the páramo, usage of chemical ferti1izinto the Independence period, agriculture and cattle- ers, insecticides, pesticides, and fungicides increased raising became common (Velázquez, 1986) and the (Bemsen, 1991; Ferwerda, 1987; López-Zent, 1993), range ofpáramo-like habitats probably expanded. followed by soil contamination. For instance, in the
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33
Fig. 20. House or "choza" at Pozo Azul on the ascent ofNevado del Cocuy (Boyacá, Colombia) at 411 Om, after a night oflight snow. Walls ofthe house are formed from the stems of Espeletia lopezii and the roofthatch from the bunchgrass Calamagrostis sp. These "bornes away from home" are often used by local inhabitants as temporary shelter during travel or periods of harvest or herding. The travelers shown in this photo are (from the right) José Cuatrecasas, H. García-Barriga, Martínez Zuloaga, Páez, and two guides. (Photo Cuattec. C-1221A taken in 1938.)
crop belt ofthe Venezuelan páramos (3000-4000 m), the traditionalland-use system of short periods of crop production (1-4 years) altemating with fallows of720 years-which allows 1ong-term recuperation of nutrients with negligible erosion rates and maintenance ofhigh 1evels of soil organic matter-has been interrupted by massive use of chemical fertilizers and a market-oriented agricultura! system. Sorne of the consequences have been "greater incidence of pest outbreaks, soil exhaustion, a possible decrease in soil organic matter, and intensification of erosion" (Sarmiento M. et al., 1990, 1993). According to Ferwerda (1987), more than 70 years are needed for regeneration of páramo bunch-grasslands after patato cultivation. Along with the Upward movement of agriculture and increased livestock production have come 1.he continued lowering ofthe forest line due to wood cutting to make more room for crop production
and pastureland (Kok et al., 1995). The timber is used (locally and regionally) for building, fence posts, fue!, etc. Fuelwood collection, mainly for cooking purposes, also removes shrubby vegetation from the páramo proper (Becker, 1988). Verweij (1995) estimated that an average family of seven persons uses about 20 kg offuelwood every day, which corresponds to a cleared area of one hectare per year. Other recent developments that have led to additional páramo disturbance include construction of aqueducts, drainage systems, and roads ( Young, 1994), an increase in the number of off-road vehicles (Pérez, 1991 b), mining (especially of sand, grave!, and peat), and afforestation projects (Rangel Ch., 1989; pers. observ.). The impact ofplantations ofexotic tree species (especially Pinus radiata and P. patula) on the development of soils and vegetation placed in or near (extensively grazed) páramo has recently been
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MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
described by Hofstede (1997). Even tourists are now becoming a problem in sorne areas (Horn, 1986; Urgilés Sánchez, 1990).
Impact of Burning and Grazing Fire is used in the páramo for two main purposes. First, it is a means to clear the upper limits of forest, thus providing open grasslands for grazing oflivestock and new land for cultivation of crops. Second, because forage quality is poor and up to 80% of the aboveground biomass ofthe tussock- or bunchgrass growth form is dead material (Cardozo & Schnetter, 1976), farmers bum offthe old or dead nonpalatable parts to provide new, more nutrient-rich growth for cattle. Ramsay and Oxley ( 1997) say that it is quite difficult to estimate the ultimate effects offire on páramo vegetation and its species diversity, since fire has been so mucha part ofpáramo history. Nevertheless, buming and grazing go hand-in-hand in most high-elevation areas of the northem Andes; seemingly they are the only applied "management practice." However, in view of the fact that it has be en part of páramo history sin ce at least Holocene times, is fire necessarily bad for páramo? And what are the actual effects of fire and grazing on páramo? How do the continuous and repetitive practices ofbuming and grazing combine to affect species composition, nutrient and soilwater content, biomass and productivity, decomposition, and short-term and long-term stability of the páramo ecosystem? The following discussion out! in es what is currently known about the role and impact of fire on the páramo ecosystem. Four recent studies about the combined effects of buming and grazing on the páramo ecosystem contribute new insight and provide valuable basic information that is necessary for future management programs (Hofstede, 1995c; Keating, 1995; Ramsay, 1992; Verweij, 1995). In the following section, I ha ve drawn heavily from the excellent dissertation ofRobert Hofstede ( 1995c ), who studied the ecological impacts ofburning and grazing on biomass, nutrient status ofvegetation and soil, and hydrology, in the Los Nevados National Park (Cordillera Central, Colombia). In a complementary study, Pita Verweij (1995) focused on the impacts ofmanagement on vegetation development and landscape ecology in the same national parle. In addition, Paul Ramsay (1992) and Philip Keating (1995) studied páramo vegetation in Ecuador, including community ecology, dynamics, and productivity. Sorne oftheir main conclusions are summarized in the following paragraphs. Hofstede (1995c), Lregaard (1992), Ramsay (1992), and Verweij (1995) all conclude that natural
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páramo vegetation is to a certain extent tolerant toward the management practices ofbuming and grazing, and that fire does offer sorne short-term advantages to the cattle-farmer (overall palatability, initial productivity). It is not, however, a wise long-term management too!, because when the burning frequency is more than once per decade (Verweij, 1995) and grazing intensity increases, then stress on the vegetation is too great, effects are mostly negative, and there is degradation ofthe ecosystem, including a loss in species diversity. Unfortunately, it has been shown that farmers will choose the short-term solution-i.e., rapid post-fire regeneration of palatable forage for maximum livestock production-at the expense of · long-term degradation of the vegetation and soil (Hofstede, 1995; Ramsay, 1992; Verweij, 1995). Burning and grazing affect the páramo ecosystem most importantly in the following three ways: 1) vegetation structure, composition, and species diversity; 2) soil structure and water-storage capacity; and 3) decomposition, nutrients, and productivity. VEGETATION STRUCTURE, COMPOSITION, AND SPECIES DIYERSITY
Despite their many adaptations, rosette plants (especial! y members of the Espeletiinae) and tussocks or bunchgrasses are sorne of the most susceptible species to burning and grazing in the páramo. In Venezuela, Smith (1981) found that 55% of the adult Espeletia schultzii (Asteraceae: Espeletiinae) plants could be killed by fire in a dry season, while in Colombia, Verweij and Kok (1992, 1995) found greater than 80% mortality in juveniles of E. hartwegiana immediately after fire. Verweij and Kok (1992) also found that grazing hada major impact on adult mortality of E. hartwegiana in Colombia, and Pérez (1992a) found that cattle browsing c.ould cause a decrease of 37% in rosette cover of Coespeletia timotensis (Asteraceae: Espeletiinae) in Venezuela. In southemEcuador, Keating (1995) found thatburning and cutting reduced the number of rare species and the relative frequency of shrub species. In the Buenavista subpáramos of Costa Rica, Horn (1997, in press) found that 90-99% ofthe dominant bamboo, Chusquea subtessellata, and the ericaceous shrub Vaccinium consanguineum resprouted following fire, whereas the second most common woody species, the shrub Hypericum irazuense (Clusiaceae), súffered 94% mortality. This display of fire sensitivity in woody species certainly affects species composition and may be the reason for the present dominance of the bamboo. For tussocks and bunchgrasses, buming and subsequent grazing also lead to fragmentation be-
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cause of trampling by livestock after the fire. Continued grazing 'or overgrazing leads to replacement of the tussocks and bunchgrasses by short-grass communities (Agrostis, Festuca, Paspalum), rosaceous species (Acaena and Lachemilla orbiculata), and species of low or matted herbs such as Eryngium humile (Apiaceae); Bidens, and Hypochaeris (Asteraceae) (Fig. 10B). These plants are better adapted to cattlegrazing and to more exposure of bare ground. Alien species, which are frequently found in these communities, are indicators ofhuman and grazing (with sorne fire) disturbance; these in elude (among others) Achillea millefolium, Gamochaeta americanum (formerly Gnaphalium americanum), Stevia lucida, and Taraxacum officinale (Asteraceae), Arenaría serpens, Cerastium arvense, and Stellaria media (Caryophyllaceae), Trifolium repens and Lupinus spp. (Fabaceae), Geranium multipartitum (Geraniaceae), Juncus effusus (Juncaceae ), Oxalis spp. (Oxalidaceae), Anthoxanthum odoratum, Dactylis glomerata, Digitalis purpurea, Poa annua, and Holcus lana tus (Poaceae ), Rumex acetosella (Polygonaceae ), and the fem Pteridium aquilinum (Dennstaedtiaceae) (Cleef, 1981b; Cleef & Rangel Ch., 1984; Ewe1 et al., 1976; Hofstede, 1997; Hofstede, Chilito P. & Sandoval S., 1995a, 1995b; Keating, 1995; Ramsay, 1992; Rangel Ch., 1989; Salamanca, 1991; Smith, 1974a; Verweij & Budde, 1992; Verweij & Kok, 1995). Based on studies of small-scale dynamics within severa! páramos in Ecuador, Ramsay (1992) generalized that a) it is mainly vegetation structure that determines fire temperature, with the highest temperatures (to 600°C) produced in the tussock canopy and the lowest (ca. 65°C) found within the tussock base and just beneath the soil surface; b) the main form of recovery from fire is regeneration by vegetative reproduction ofbelow-ground plant parts; e) the degree of survival of intertussock species depends on the severity ofthe fire, which is largely a function ofthe time interval since the last burning; d) survival following fire does not guarantee a plant's persistence in the community, since both tussock grasses and intertussock species show significant mortality rates in the months following fire; e) recovery at higher altitudes is slower; t) burning may cause the pattems of community development to become cyclical; g) sorne species that quickly colonize bare ground by seed may persist, but as the vegetation matures most are killed by competition; and h) establishment within tussocks after burning is the preferred method of sorne species, such as Lupinus cf. pubescens. Minimum fire recovery periods, i.e., the period of post-fire growth required to generate enough fue! to carry a subsequent fire (Verweij, 1995), of3-10 years
35
have been reported depending on the vegetation structure (Horn, 1990a, i991; Janzen, 1973; Ramsay, 1992; Ramsay & Oxley, 1996; van Groen, 1987; Verweij, 1995). However, the frequency ofburning is typically every 2-4 years (Ramsay & Oxley, 1996), and in practice, fire is applied too frequently to allow full bunchgrass recovery (Verweij, 1995). Minimum fire recovery periods do not mean that the vegetation will ha ve recovered completely, however, because species composition and distribution will change and more bare spots will be present (Hofstede, 1995c; Horn, 1989a; Janzen, 1973; Miller & Silander, 1991; Verweij, 1995; Williamson et al., 1986). Furthermore, since grasses have high relative growth rates, and burning and grazing stimulate tillering, they are favored over woody species by high fire frequencies (Horn, 1991; Janzen, 1973; Ramsay, 1992; Ramsay & Oxley, 1996; Verweij, 1995; Williamson et al., 1986). Verweij (1995) concluded that more than 1O years are needed before the average number of vascular plant taxa stabilizes back to pre-fire levels, implying also that fire frequency may influence species richness in the vegetation. In summary, it would appear that occasional disturbances over short intervals may allow for relatively rapid recovery of community structure and composition, but that repeated burning, combined burning and grazing, or severe disturbances at intervals of even 24 years are too great to allow complete recovery of páramo vegetation and sorne species are lost. SO!L STRUCTURE AND WATER STORAGE CAPAC!TY
The páramo belt is an indispensable source ofwater for drinking and irrigation for a majority of the people in Venezuela, Colombia, and Ecuador. In its natural, undisturbed condition, páramo acts as a sponge, where the excess water is slowly but constantly returned to the ecosystems oflower elevations (GómezMolina & Little, 1981). Natural páramo shrub vegetation holds up to 12 times more water than a disturbed grass community (Schnetter et al., 1976), while water retention in soils of the páramo surpasses 200% of its own dry weight (Cañadas Cruz, 1983). Hofstede and Sevink (1995) showed that undisturbed páramo has a larger water-storage capacity than bumed and grazed páramo, and that a continuously wet plant !ayer is important for maintaining ihe large soil storage for drier periods. With grazing and burning, however, vegetation cover disappears and is replaced by a low-growing ground cover that cannot hold as much moisture since the páramo soils are more compressed, drier, less acidic, and somewhat less organic (Hofstede, 1995c). Verweij and Budde (1992) and Verweij (1995) also found that during the first
36
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
1.5-5 years following fire the percentage ofbare soil increased by more than 10%, depending on the vegetation structure and grazing intensity. Meanwhile, vegetation is trampled and compacted and the turf is broken-up by cattle (Grubb, 1970; Pérez, 1993a), thus favoring eros ion, loss of soil, and micro-terracing, especially on slopes (Flores Ochoa, 1979; Pérez, 1992b, in press; Troll, 1973). DECOMPOSITION, NUTRIENTS, AND PRODUCTIYITY
Hofstede (1995c) and Verweij (1995) have shown that grazing and especial! y buming practices result in faster decomposition and mineralization dueto drier soils and higher maximum soil temperatures, but this does not necessarily lead to higher nutrient contents of soils and plant tissues. On a short-term basis, mineralized nutrients may become available for imq¡ediate plant regrowth, but generally they are very quickly unavailable in the soil due to immobilization especially of phosphorus, volatilization especial! y of nitrogen and sulphur, or, less important, erosion or leaching. Therefore, the vegetation remains nutrient poor. The change in vegetation structure and composition eventually leads to less tussocks, which may have more tillers per unit area anda higher Ji ve material content, but the overalllower stature and leaf elongation rates yield less productivity (Hofstede, 1995c ). This low productivity, combined with nutrient-poor vegetation, results in poor-quality cattle (Hofstede, 1995d; Pels & Verweij, 1992; Schmidt & Verweij, 1992; Verweij, 1995). In summary, the general impact ofincreased buming and grazing was summarized by Verweij (1995) as follows: The structure ofthe bunch-grasslands becomes more open. Fire events cause temporal changes in vegetation structure o ver the relatively short time span of a few years. Without the influence of grazing, the vegetation is able to recover and retums to the initial structure of dense bunchgrassland, the floristic composition remaining largely unchanged. Under prolonged grazing pressure, however, short matted grasses and forbs increase anda gradual transformation into other communities occurs. Changes in structure are of vital importan ce in relation to the water holding capacity of both vegetatiorí and soil. Hofstede & Sevink (1995) concluded that water retention capacity is reduced in grazed and burned situations, with an increased risk of drying especially in the drier seasons.
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The Fu tu re of the Páramo Ecosystem The páramo is rapidly becoming a threatened ecosystem. With human population size at its greatest in history and steadily rising, pressure for greater food production and land use has intensified and deterioration of the environment has inevitably followed. Activities such as deforestation, road building, buming, agriculture, and herding of animals ha ve significantly altered this fragile ecosystem. They have also caused native páramo plants and timberline forests to disappear atan alarming rate, thereby allowing weedy species to domínate sorne landscapes. Such activities also Iead to accelerated erosion on the slopes and flooding;· as the soils are notable to retain moisture within tlieir drainage basins. Furthermore, with the publi~:'s increased desire for leisure time anda gradual awakeriing to the potentia! of the páramo for recreation and tourism, we need to quickly increase our efforts to study this ecosystem, not only to know what is there by means of inventory and ecological analyses, but also to monitor the effects of disturbance on the vegetation and aquatic resources so that rational management programs can be established to protect and preserve the páramo. Why is the páramo important and what needs to be done to maintain this ecosystem? Can we still turn the ti de of alteration and destruction? Must it be totally preserved and protected? As has been discussed above, páramos are important for scientific, ecological, and economic reasons-all ofwhich are interconnected and interdependent. Sorne ofthese reasons are summarized below, and opinions are offered about what should be done. SCIENTIFIC REASONS
Páramos occupy no more than 2% ofthe land area of the northem Andes in Venezuela, Colombia, and Ecuador, and yet they are highly di verse biologically and contain many plants that grow nowhere else in the world. Their geographical isolation along the mountainous chain from Costa Rica to northem Peru and their high diversity and endemism combine to malee them biogeographically unique. Paleoecological studies indica te that páramo habitats have undergone repeated expansion and contraction during the late Pliocene and Pleistocene. These high-elevation, cold-climate ecological islands of endemism are surrounded by a sea oflow-elevation, warm tropical forest. Therefore, the insular dispersa!, age, origin, affinities, and evolutionary pattems ofthe plant taxa are open to detailed systematic and evolutionary biogeographic studies. Human activities, however, are severely limiting biodiversity in the páramo ecosystem. Basic facts
1999]
PARAMOS
about the taxonomy, relationships, distribution, and ecological preferences of most high-elevation groups is very incomplete. We must know the names ofthe taxa we are dealing with before we are able to compare them or effectively communicate data about their ecology, evolution, and so forth (Vuilleumier & Monasterio, 1986). Therefore, it is appropriate and timely that páramo plants be studied and a complete flora with keys, species descriptions, and indications of geographical distribution be written. EcoLOGICAL REASONS
Perhaps the single most important ecological function ofpáramo is as a regulator of Andean hydrology. However, deterioration of the páramo due to loss of vegetation cover, soil compaction, and weakened water storage capacity may increase soilloss, erosion, ard flooding and contribute to droughts during dry periods. Therefore, ecological studies must also begin at the basic leve!, including additional short- and longterm studies ofthe dynamics ofpáramo vegetation as it affects landscape heterogeneity and plant species diversity, keeping in mind the use ofbetter management techniques that could lead to stabilization of water supplies, increased productivity, and a subsequent lesser need to cut more forest and exploit new areas of land (Ramsay, 1992; Verweij, 1995). Also needed is a better understanding of the dynamics of fire, repeated fires, and recovery after fire in order to properly assess its role as a too! in the management and conservation of páramo ecosystems. It has also been noted that the páramo is a very sensitive ecosystem and could provide an excellent area for additional environmental studies of global and regional climatic changes, such as the effects of global climate change (the greenhouse effect) and drought or increased rain (brought on by El Niño) on plant species distributions and adaptations. EcoNOMIC REAsONS
The major economic use of the páramo toda y is for agriculture and livestock. But, as has been discussed abo ve, the management practices of buming and grazing, and the increased use of chemicals, emphasize only short-tem1, not long-term, usage. Ifthe páramo ecosystem is going to remain an important ecological and economic center, it must continue to be the natural source of clean water as well as a resource for agriculture and grazing, new foods, ornamentals and medicines, and recreation and tourism. Páramo plants are contributors of important genetic factors to high-elevation or cool-habitat crops such as
37
the potato (Solanum tuberosum, Solanaceae) and the grain quinoa (Chenopodium quinoa, Chenopodiaceae ). Páramos are also a source of potential new foods ("new" to non-Andean peoples) derived from pre-Columbian tuber crops such as "oca" or "íbias" (Oxalis tuberosa, Oxalidaceae), "ulluco" or "olluco" (Ullucus tuberosus, Basellaceae), and "añu," "mashua,".or "cúbios" (Tropaeolum tuberosum, Tropaeolaceae) (King, 1988; National Research Council, 1989; Schjellerup, 1989). New ornamental plants such as Senecio niveo-aureus (Asteraceae), and new medicines such as those derived from Oritrophium spp. and Chuquiraga jussieui (Asteraceae) and Thamnolia verm icularis (a lichen of uncertain affinity), have as yet unrealized potential (Aranguren B. & Márquez, 1995; Aranguren B. et al., 1996; López-Zent, 1993; Paredes, 1962; Pérez, in press). Historically, the páramo has been a source of traditional medicines for local communities, and even today 56% (43 out of77) ofthe medicinal plants used by the Cogui, an indigenous people living in the Sierra Nevada de Santa Marta (Colombia), come from the páramo (Carbonó de la Hoz, 1987). Although there are limitations, the potential for ecotourism or for recreation-fishing, hiking, camping-in páramo regions has gone almost unnoticed (Black, 1983, 1988; Ramsay, 1992; Rangel Ch., 1989; Urgilés Sánchez, 1990). The so-called wilderness experience can draw many visitors, but, as Hofstede (1995c) has reminded >IS, these same visitors are disappointed when meeting unnatural elements such as cattle, fences and meadows. Both national and intemational tourists potentially visiting the páramo are not directly interested in a high species richness, a high water storage or a good soil protection, but their demand is that the landscape is as original as possible: they simply enjoy touching the snow and seeing an Espeletia. Nonethe1ess, the ecological and hydrological sustainability benefits directly from the conservation of an original landscape and vegetation structure .... In the future, it should be evaluated which yields more money: a productive cow ora majestic Frailejón. What can be done? Virtually any extended use ofthe high-elevation páramos-be it for science, economics (including both resource production and extraction), or conservation-will, to sorne extent, have an impact on species composition, vegetation structure, biodiversity, and soil hydrology. It is only if all human activities in páramos are ceased that ecological
38
MEMOIRS OF THE NEW YORK BOTANICAL O ARDEN
and hydrological sustalnability will be assured. With the current socioeconomic situation in the Andes, however, the exclusion of all human activities from the páramo is not realistic, and agricultura! production has to be accepted and respected (Hofstede, 1995c; Lauer, 1993; Verweij, 1995). Given the scientific evidence at hand, it is relatively easy to identify the major problems that have changed the páramo ecosystem-but what are the practica! alternatives? At this time, three areas in particular need to looked at: management practices, conservation efforts, and environmental education. MANAGEMENT
It seems generally agreed upon that there is not any single management practice that addresses all páramo functions; therefore, perhaps the challenge to managers is to vary their programs in order to meet the demands of one function in one area and those of another somewhere else (Hofstede, 1995c; Horn, in press; Pérez, in press; Ramsay, 1992; Verweij, 1995; Young, 1997). While banning the use ofburning is advocated by sorne (e.g., Hofstede, 1995c; Verweij, 1995), Horn (in press) believes that fire management efforts need to be changed from the prevention of all fires to the control of fire frequency, thus recognizing its place in the páramo landscape. Ramsay ( 1992) advocated shifting agricuiture and grazing to lowerelevation zones, since it appeared from the results of his productivity studies that lower-elevation pastures were "silbstantially underproductive." Rangel Ch. (1989) suggested severa! measures: declaring all páramos as nationally protected areas, primarily to protect the water resources; restricting use of all páramo bodies ofwater, especially in the production of energy; avoiding the exploitation of páramo biological resources and especially avoiding grazing by sheep; establishing research centers to study páramo; and using particular páramo areas for recreation (with controls). Numerous smaller, but nonetheless important, changes should also be considered for implementation wherever possible: prohibiting the cutting of natural vegetation and the collection of fallen stems for fue!, creating fenced enclosures to exclude cattle or allow rotation of grazing, and excluding motorcycles and off-road vehicles from the páramo (Pérez, in press). If new management strategies are to be considered, there needs to be strong coordination and cooperation between scientists, local people, nongovernmental institutions, and government departments in the entire process ofplanning, design, and implementation.
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Becker (1988) has stressed that within indigenous communities there are needs for alternative sources of income to alleviate the pressures ofhabitat destruction for agricultura! purposes (see al so Verweij et al., in press), better social organization with the promotion of traditiona1 techno1ogies such as terracing for soil conservation (Lauer, 1993; Nieto Cabrera et al., 1997; Sarmiento M. et al., 1993), reforestation with useful and native species (Brandbyge, 1984; Brandbyge & Holm-Nielsen, 1986; Lauer, 1993; Sarmiento M. et al., 1993), and integrated agroforestry systems and controlled grazing and burning to allow regeneration of native plants (Hofstede, 1995c; Ramsay & Oxley, 1996; Verweij, 1995). CONSERVATION
Because oftheir high biological diversity and their biogeographica1, evolutionary, ecological, and economic significance, páramos are unique and the species that have adapted to them will not grow elsewhere. These facts, along with the realization that the páramo, like so many other tropical ecosystems, is being altered by human activities, qualify them as high-priority areas for conservation. "Their preservation in the face of these ever-expanding human activities, however, will depend on their rational use" (Vuilleumier & Monasterio, 1986). The farmer's aim in buming is to increase livestock productivity, a practice that may work in the short-term; however, the process of recovery in páramo takes a long time and subsequent repeated burning and grazing may cause long-term damage, in complete contrast to the original aim of the burning (Ramsay, 1992). Therefore, conservation of the páramo ecosystem, including watersheds and natural vegetation, is imperative. Until about 40 years ago there was little direct action taken toward preservation of the páramo ecosystem. Recently, however, páramos have been included within the boundaries of many national parks, frequently with recognition as regulators of Andean hydrology. National parks that include important stretches ofpáramo are Chirripó (Costa Rica); Sierra Nevada de Mérida and Tamá (Venezuela); Sierra Nevada de Santa Marta, Sierra Nevada de Cocuy, Sumapaz, Chingaza, Los Nevados, and Puracé (Colombia); Cotopaxi, Sangay, and Podocarpus (Ecuador); and the Río Abiseo watershed in northern Peru. More representa ti ve páramo areas still need to be set aside, however, and there must be greater efforts to increase the numbers of rangers to enforce existing boundaries and park regulations.
1999]
PARAMOS
ENVIRONMENTAL EDUCATION
Scientists have the responsibility to see that factual data from páramo studies be presented to the general public, local authorities, and indigenous communities, because public awareness and public support are essential to the achievement of conservation ancl/or management goals. People from large cities like Bogotá, Quito, Cuenca, and Mérida flock to the páramos on weekends for the natural beauty, the "wilderness experience," and recreation, but at the same time they often use branches from shrubs for their campfires and leave behind their litter (Urgilés Sánchez, 1990). We must make the general public aware of the value of páramo, which can also serve as an educational model in student field trips and ecological tours. For instance, one excellent means of educating the public about dif-
39
ferent aspects of the environment is to place narrative signs along roads throughout the parks. If, as Horn (in press) advocates, fire is recognized as having a place in the páramo landscape, then park managers will also need to educate the general public about the role of fire in maintaining diversity ofhabitats and species. The training oflocal students is also critica! for longterm maintenance of diversity and identification of critica! areas for protection and management (Balslev & Paz y Miño C., 1991; Verweij, 1995). For logistical reasons, local scientists can more easily undertake long-term surveys and their follow-ups. Furtherrnore, many young scientists will become key government officials or leaders in conservation groups in their countries, thus improving the quality of decisionmaking about conservation and management issues.
CHECKLIST
OF
PARAMO
The following checklists for lichens, mosses, hepatics, and vascular plants were written by and are the responsibility of their respective authors. Each section begins with a brief introduction stating the concept of páramo used by each author and providing general information about the group. This is followed by a species list compiled by each author, based on his concept of páramo as stated in his introduction. The checklists are arranged alphabetically by family and, within each family, by genus and species. The first family and/or genus entry may be followed by one or more literature references. Each specific entry gives the proper genus and species name, authority, relevant synonymy, geographical distribution of the species, elevational range of the species, and sornetimes brief statements conceming inclusive taxa, origin, use, etc. Fom1al notes are referred to by either the word "Note(s)" or by numbers in square brackets; normally, they are listed immediately after the entry (family, genus, or species names) to which they refer, or rarely at the end of the entry to which they refer. A question mark (?) after an entry s_ignifies the
PLANTS
author's doubt asto presence of a taxon in the country (e.g., EC?), or sorne degree of uncertainty about exact elevation (e.g., 3500-? m). Finally, a short bibliography specific to that group of plants is given, usually referring only to papers cited in the text and generally taxonomic in nature; these usually are not repeated in the overall páramo bibliography at the end of the book. Herbarium acronyms follow the most recent edition of Index Herbariorum (Holmgren et al., 1990). Author abbreviations follow Authors ofPlant N ames (Brummitt & Powell, 1992). Symbols used in the checklists, referring to the geographical range of páramo countries, are given in the following sequence: CR =Costa Rica, PA= Panama, CO = Colombia, VE =Venezuela, EC = Ecuador, and PE = northem Peru. Literature Cited Brummitt, R. K. & C. E. Powell (eds.). 1992. Authors of plant names. Royal Botanic Gardens, Kew. Holmgren, P. K., N. H. Holmgren & L. C. Barnett (eds.). 1990. lndex herbariorum. Part !: The herbaria of the world. Ed. 8. Regn. Veg. 120. International Association for Plant Taxonomy, Bronx, NY.
Lichens H. J. M. Sipman The species listed reflect this preference for azonal habitats: a considerable group inhabits xerothermic habitats and has its main distribution at lower elevations, including species of Catapyrenium, Toninia, and Xanthoparmelia. The same holds true for most species from tree trunks, which equally seem to be outposts of taxa from lower elevations. The genus Stereocaulon, on the contrary, is clearly more diverse in the páramos, probably because it grows on free rock faces. At lower elevations it can be found mainly where human activities have created suitable habitats in road cuts or in deforested areas. The checklist is based principally on an evaluation ofthe literature. The main sources were a list ofmacrolichens from parts of Venezuela (López-Figueiras, 1986), a survey of lichen zonation in Ecuador
This checklist presents all species of lichenized fungi currently known from the páramo region and includes 45 families, 114 genera, and ca. 463 species. Neither subspecific taxa or lichenico1ous fungi (fungi that grow on lichens) are included in the list. For the preparation of the checklist, the páramo region is defined as the area above 3200 m elevation in Costa Rica, Panama, Venezuela, Colombia, Ecuador, and northernmost Peru. Vegetation characteristics have not been used for delimitation, because literature references and herbarium labels tend to give little information on the vegetation. Also, lichens depend on the availability ofmicrohabitats, such as rock outcrops and tree bark, rather than on the zonal vegetation. For example, pure grassland, including páramo grassland, tends to be poor in lichens. 41
42
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
(Arvidsson, 1991), a checklist for the Venezue1an páramos (Marcano et al., 1996), and, in particular, the information accumulated for a planned checklist of Co1ombian lichens. For completion, information was taken from herbarium specimens archived at B, and to a lesser extent at NY, U, and in the herb. Aptroot. However, the evaluation of the herbarium material was not complete, and various additions can be expected, in particular from NY, GB, and US. Early literature required special attention, because different species concepts were sometimes used. Where an occurrence in the páramos seemed unlikely in view ofthe presently known distribution, it is supposed that the recorded material does not belong to the species in its current sense, and the record has been omitted. However, the actual distribution is poorly known for many species, so that sorne doubtful records may still be listed. Checking of specimens, on which the records are based, has not been done. Literature references are given at the beginning of each family or genus. For species where no published information was available, a single representative . specimen is indicated. The indicated altitudinal range concems only the countries where páramos are found. In other countries the altitudinal range of a species may be considerably different from that listed here. Many species have large ranges, extending far beyond the páramos, and may occur at considerably lower elevation in, e.g., temperate areas with an oceanic climate. This list is probably more incomplete than those of the other groups in this book because of a lack of distribution data and because of incomplete taxonomic knowledge of lichens in general. Few páramo areas have been investigated more than superficially for their lichen flora, and the results ofthese explorations are only to a small extent published. In sorne publications, no precise indication of elevation is given, so that páramo records could not be recognized as such. Few groups have received sufficient taxonomic attention to make reliable determinations possible. Among the fairly well-known groups are the families Cladoniaceae and Parmeliaceae (excl. Usnea), and the genera Stereocaulon and Umbilicaria. An increased knowledge ofpáramo lichens will certainly show that sorne ofthe poorly known genera are much more diverse in the páramos than the present list suggests. Furthermore, it seems probable that at least twice as many species will eventually be found and that most taxa will eventually be found in all páramo countries. Lichens have wide geographical and altitudinal ranges, as compared with seed plants. Consequently, most species extend far beyond the páramos, and few páramo endemics are expected. For an analysis ofthe geographical affinities of the páramo lichen flora, se e
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Sipman, 1992b. Nevertheless, there are indications for floristic differentiation within the páramo area. The southern part, from southern Colombia southward, seems to have a markedly different lichen flora, as could be observed in the genera Everniastntm (Sipman, 1986a), Stereocaulon (Sipman, 1987), and Umbilicaria (Sipman, 1992a). A restricted range in the southern part is fairly well documented for, e.g., Cetrariastntm ecuadoriense, Stereocaulon globuliferum, and S. novogranatense. A well-documented species apparently restricted to the northem part ofthe páramos is Cladina halei (Ahti, 1986). Table II lists the 10 largest families and genera of páramo lichenized fungi.
CHECKLIST OF LICHENS
Lichenized Ascomycetes ACAROSPORACEAE Re f.: Hertel, 1971. Polysporina Vezda P. simplex (Davies) Vezda [syn.: Sarcogyne simplex (Davies) Nyl.]; VE; 4250 m
ARTHOPYRENIACEAE Ref.: Hawksworth, 1985. Mycomicrothelia Keissl. M. apposita (Nyl.) D.Hawksw.; CO; 3500 m
ARTHRORHAPHIDACEAE Ref.: Arvidsson, 1991; Hertel, 1971; Sipman, 1989. Arthrorhaphis Th.Fr. A. alpina (Schaer.) R.Sant. [syn.: A. citrinella var. alpina (Schaer.) Poelt]; CR, CO, VE, EC; 3500~300 m A. citrinella (Ach.) Poelt; VE, EC; 2850-4500 m
ASTEROTHYRIACEAE Asterothyrium Müll.Arg. A. leptosporum Müll.Arg.; CO; 3200 m [Note: leg. Sipman & Aguirre 27486 (B)]
BACIDIACEAE Bacidiopsora Kalb Ref.: Kalb, 1988. B. squamulosula (Nyl.) Kalb; EC; 1750-3200 m Hypocenomyce M.Choisy Ref.: Sipman, 1989. H. scalaris (Ach.) M.Choisy; CO; 3750 m
43
PARAMOS
1999]
CHRYSOTHRICHACEAE
Phy/lopsora Müll.Arg. Ref.: Brako, 1991; Kalb, 1988. P. buettneri (Müll.Arg.) Zahlbr. var. glauca (de Lesd.) Brako; VE; 1400-3500 m P. confusa Swinscow & Krog; VE; 1600-3500 m P.fendleri (Tuck. & Mont.) Müll.Arg.; VE; 2400-3600 m P. parvifolia (Pers.) Mü1l.Arg. var. parvifolia; EC; 3200 m
Chrysothrix Mont. C. chlorina (Ach.) Laundon [syn.: Lepraria chlorina (Ach.) Ach.]; CO, VE; 3500-4175 m
Tephromela M.Choisy Ref.: Hertel, 1971; Ny1ander, 1863a, 1863b. T aglaea (Sommerf.) Herte1 & Rambold; VE; 4000-4700 m T atra (Huds.) Hafellner [syn.: Lecanora atra (Huds.) Ach.]; CO, VE; 1400-3900 m T elixii Kalb;'VE; 3500 m
CladiaNyl. Ref.: Arvidsson, 1991; Filson, 1981; López-Figueiras, 1986. C. aggregata (Sw.) Nyl.; CO, VE, EC; 1300-4550 m C. fuliginosa Fi1son; CO, EC; 3605-4040 m [Note: This concerns probably an undescribed species different from the Australasian C.fuliginosa, under study by T. Ahti.]
BAEOMYCETACEAE Ref.: Arvidsson, 1991; López-Figueiras, 1986; Vries & Sipman, 1984. Baeomyces Pers. B. rufus (Huds.) Rebent.; CR, CO, VE, EC; 3350-4150 m Phyllobaeis Gierl & Ka1b P. erythre/la (Mont.) Kalb [syn.: Baeomyces imbricatus Hook. var. e~ythrellus (Mont.) de Vries]; VE, EC; 18003452 m P. imbricata (Hook.) Kalb & Gierl (syn.: Baeomyces imbricatus Hook.); CO, VE, EC; 1200-4400 m P.linearis (de Vries) V.Marcano & Sipman; VE; 2400-3400 m
CALICIACEAE Ref.: Sipman, 1989; Tibell, 1996. Calicium Pers. C. glaucellum Ach.; CO; 1750-3750 m C. salicinum Pers.; CO; 3500-3750 m C. subquercinum Asahina; CO; 3500 m
CANDELARIACEAE Candelaria A.Massal. Ref.: López-Figueiras, 1986. C.fruticans Poelt & Oberwink1er; CO, VE; 1100-3500 m Candelarie/la Müll.Arg. Ref.: Hertel, 1971. C. vi tellina (Hoffm.) Müll.Arg.; VE; 3900 m
CATILLARIACEAE [Note: The systematic position of these two genera is unc1ear; they are placed here provisionally.] Lopezaria Ka1b & Hafellner [1] Ref.: Sipman, 1989. L. versicolor (Fée) Ka1b & Hafellner [syn.: Catinaria versicolor (Fée) Sipman]; CO, VE; 300-3500 m Sporastatia A.Massal. Re f.: Hertel, 1971. S. testudínea (Ach.) A.Massal.; VE; 4250 m
Ref.: Sipman, 1989.
CLADONIACEAE
Cladina (Nyl.) Nyl. Ref.: Ahti, 1961, 1986; Arvidsson, 1991; LópezFigueiras, 1986; Sipman & Cleef, 1979. C. arcuata (Ahti) Ahti & Follmann (syn.: Cladonia arcuata Ahti); CO, VE, EC; 1900-3950 m C. boliviana (Ahti) Ahti; CR, CO, VE, EC; 1150-4000 m C. confusa (R.Sant.) Follmann & Ahti; CR, CO, VE, EC; 1280-4000 m; includes fo. bicolor (Müll.Arg.) Ahti and fo. confusa C. halei Ahti; CO, VE; 2000-4100 m C. rangiferina (L.) Nyl. subsp. abbayesii (Ahti) W.L.Cu1b. [syn.: Cladonia rangiferina (L.) F.H.Wigg. var. abbayesii Ahti]; CO, VE; 1500-4050 m Cladonia Hill ex P.Browne Re f.: Ah ti & Stenroos, 1986; Arvidsson, 1991; LópezFigueiras, 1986; Sipman, 1989; Stenroos, 1989a; Vainio, 1894, 1899b. C. aleuropoda Vain.; CO, EC; 3800-4250 m C. andesita Vain.; CR, CO, VE, EC; 2100-3900 m C. bacillaris Nyl.; VE; 1400-3600 m C. calycantha (De1ise) Nyl.; CO, EC; 2750-3700 m [Note: See Ahti & Marcelli, 1995.] C. carassensis Vain.; VE; 1800-3450 m C. cartilaginea Müll.Arg.; VE; 1200-3450 m C. chlorophaea (F!Orke ex Sommerf.) Spreng.; CO, VE, EC; 2100-4350 m C. chondrotypa Vain.; CO; 1900-3300 m [Note: leg. Aguirre & Sipman 5858 (B)] C. coccifera (L.) Willd.; CR, CO, VE, EC; 2150-4350 m C. corniculata Ahti & Kashiw.; CO, VE, EC; 2000-4350 m C. corymbites Nyl.; VE; 3500 m C. corymbosula Nyl.; CO, VE; 3750-4315 m C. crispata (Ach.) Flot.; VE; 1700-3450 m C. cryptochlorophaea Asahina; CO; 3100-38 iO m [Note: !eg. Cleef6792 (U)] C. dactylota Tuck.; CO, VE, EC; 1000-3400 m C. didyma (Fée) Vain.; CR, CO, VE, EC; 230-4050 m; ineludes var. vulcanica (Zoll. & Moritzi) Vain and var. didyma C. dilleniana Fl6rke; VE, EC;? m [Note: VE collection leg. Sipman 11225 (U)] C. furcata (Huds.) Schrad.; CO, VE, EC; 1800-4100 m
44
MEMOIRS OF THE NEW YORK BOT ANICAL O ARDEN
C. granulosa (Vain.) Ahti; VE; 1800-4000 m C. grayi Merr. ex Sandst.; CO, VE; 2300-3620 m [Note: 1eg. Aguirre & Sipman 5722 (B)] C. humilis (With.) J.R.Laundon; EC; 3400 m [Note: leg. Harris 17057 (NY)] C. isabel/ina Vain.; CR, CO, VE; 2450-4330 m C. leprocephala Ahti & S.Stenroos; CO, VE, EC; 3000-4250 m C. lopezii S.Stenroos; CO, VE; 2400-3900 m [Note: Cited as Cladonia minia/a G.Meyer in Ah ti & Stenroos, 1986 (230).] C. macilenta Hoffm. (syn.: C. bacil/aris Nyl.); CO; 12003700m C. macrophyllodes N yl.; EC C. meridensis Ahti & S.Stenroos; CO, VE, EC; 2400-4475 m C. merochlorophaea Asahina; CO, VE; 2400-3400 m; in- . eludes var. merochlorophaea in VE and var. novochlorophaea Sipman in CO [Notes: leg. Cleef9749b (U)] C. mexicana Vain.; CR, VE, EC; 2000-3500 m C. microscypha Ahti & S.Stenroos; CO, VE; 2300-3800 m C. ochrochlora Fl<'irke; CO, VE; 2000-3800 m C. pleura/a (Fiiirke) Sehaer.; CO, VE; 2900-4000 m C. pyxidata (L.) Hoffm. [syn.: C. pocil/um (A eh.) O.J.Rieh.]; CO, VE, EC; 2300-4425 m C. rappii A.Evans var. exiliar (Abbayes) Ahti; CO, VE; 1450-3800 m C. squamosa (Seop.) Hoffm.; CO, VE, EC; 1400-4510 m; includes var. subsquamosa (Nyl. ex Leight.) Vain. and var. squamosa C. subradiata (Vain.) Sandst.; CR, CO; 1900-4550 m C. subsquamosa Kremp.; CO, VE, EC; 1000-4150 m C. verruculosa (Vain.) Ahti; CO, VE, EC; 2200-4550 m
COCCOCARPIACEAE Ref.: Arvidsson, 1982; López-Figueiras, 1986. Coccocarpia Pers. C. e¡ythroxyli (Spreng.) Swinseow & Krog; CR, CO, VE; 240-3750 m C. palmicola (Spreng.) Arv. & D.J.Galloway; CR, CO, VE; 200-3500 m
[VOL. 84
L. burnetiae Dodge; VE; 2400-3500 m L. caperatum P.M.Jorg. & Wallaee; CO, VE, EC; 2000-4000 m L. chloromelum (Sw.) Nyl.; VE; 2400-3500 m L. coralloideum (G.Mey. & F1ot.) Vain.; CR, CO; 17503975 m L. cyanescens (Rabenh.) Korb.; CR, VE; 500-3500 m [Note: VE voucher 1eg. Sipman 11330c (U)] L. denticulatum Nyl.; VE; 1400-3500 m L. diaphanum (Sw.) Mont. (syn.: L. punctulatum Ny l.); VE; 1400-3400 m L. digitatum (A.Mass.) Zahlbr.; CO, VE; 1500-3975 m L.jloridanum Sierk; VE; 3750 m L. hibernicum M.E.Mitch.; VE; 2400-3450 m L. inversum P.M.Jorg. & Wallace; CO; 4070 m L. isidiosel/um (Riddle) Sierk; VE; 600-3300 m L. )Úressianum Tav.; CR, CO; 3300 m [Note: CO voucher 1eg. Aguirre & Sipman 5974b (B)] L. laceroides de Lesd.; CR, CO, VE, EC; 2000-3975 m L. mandonii P.M.Jorg.; CR, CO, VE; 2400-4325 m L. olivaceum (Hook.) Zah1br. (syn.: L.foveolatum Nyl.); CR, CO, VE, EC; 1250-3750 m L. papillosum (de Lesd.) Dodge; CO, VE; 2500-3500 m L. phy/locarpum (Pers.) Ny l.; CO, VE, EC; 1100-4600 m [Note: See Müller Argovensis, 1879: 163.] L. resupinans Nyl.; CR, CO, VE; 3000-4285 m L. stipitatum Vain.; VE; 1400-3300 m
CONIOCYBACEAE Ref.: Sipman, 1989; Tibell, 1996. Chaenotheca Zukal C. brunneola (Ach.) Mü11.Arg.; CO; 3500-3750 m C. carthusiae (Harrn.) Lettau; CO; 3500 m C. chrysocephala (Turner ex Ach.) Th.Fr.; VE; 4000 m C. gracillima (Vain.) Tibell; CO; 3500 m C. hispidula (Ach.) Zahlbr.; CO; 3500 m C. laevigata Nádv.; CO; 3750 m C. trichia/is (Ach.) Th.Fr.: CO; 3500 m
ECTOLECHIACEAE COLLEMATACEAE Ca/lema F.H.Wigg. Ref.: Dege1ius, 1974; López-Figueiras, 1986; Sipman, 1989. C. glaucophthalmum Nyl.; CO, VE, EC; 1100-4000 m; ineludes var. glaucophthalmum and var. implicatum (Ny l.) De gel.
Ref.: Santesson, 1952. Tapel/aria Mü11.Arg. T. epiphylla (Müll.Arg.) R.Sant.; CO; 1800-3200 m T. nigrata (Müii.Arg.) R. San t.; CO; 3200-3200 m [Note: 1eg. Aguirre & Sipman 5146 (B)]
FUSCIDEACEAE Ref.: Kalb, 1988.
Leptogium (Aeh.) Gray Ref.: Jorgensen, 1997; López-Figueiras, 1986; Sipman, 1989. L. aciculare P.M.Jorg.; VE; 3500 m L. andinum P.M.Jorg.; CO, VE, EC; 2000-4050 m L. austroamericanum (Malme) Dodge; VE; 3500 m [Note: 1eg. Sipman 11330b (U)] L. azureum (Sw.) Mont.; VE, EC; 1300-3500 m L. burgessii (L.) Mont.; CR, CO, VE, 2000-4325 m
Mm·onea A.Massal. M. constans (N y l.) Hepp; EC; 3200 m
GOMPHILLACEAE Echinoplaca Fée E. lucernifera Ka1b & Vezda; CO; 3200 m [Note: leg. Aguirre & Sipman 5146b (B)]
1999]
PARAMOS
Gomphillus Nyl. Ref.: Kalb, 1988. G. ophiosporus Kalb & Vezda; EC; 3400 m Gyalideopsis Vezda Ref.: Kalb, 1990. G. monospora Kalb & Vezda; VE; 3500 m G. peruviana Merr. ex Vezda; CR, VE; 3500 m
GYALECTACEAE
45
Lecide/la Kiirb. Ref.: Hertel, 1971. L. carpathica Kiirb.; VE; 4150 m L. e/aeochroma (Ach.) M.Choisy; VE; 3500-4720 m L. inamoena (Müii.Arg.) Hertel; VE; 4250 m L. /actea/a (Nyl.) Hertel & Leuckert; VE; 3500 m L. stigmatea (Ach.) Hertel & Leuckert; VE; 4250 m L. wulfenii (Hepp) Kiirb.; VE; 4150-4200 m Rhizop/aca Zopf R. me/anophthalma (DC.) Leuckert & Poelt; VE; 4000-4200 m
Ref.: Kalb, 1988. Dimere/la Trevis. D. chiodecton;ides Kalb; EC; 3200 m
HYMENELIACEAE Megaspora (Ciauzade & CI.Roux) Hafellner & V.Wirth Ref.: Arvidsson, 1991; Hertel, 1971; Sipman, 1989. M. verrucosa (Ach.) Hafellner & V.Wirth [syn.: Pachyospora verrucosa (Ach.) A.Massal.]; CO, VE, EC; 38304540m Tremolecia M.Choisy Re f.: Hertel, 1971. T. atrata (Ach.) Hertel; VE; 3900-4730 m
ICMADOPHILACEAE Dibaeis Clem. Ref.: Gierl & Kalb, 1993; López-Figueiras, 1986; Vries & Sipman, 1984. D. absolutus (Tuck.) Kalb & Gierl (syn.: Baeomyces abso/utus Tuck.); CO, VE; 1200-3 750 m D. columbina (Vain.) Kalb & Gierl; CR, CO, EC; 1200-3950 m D.fungoides (Sw.) Kalb & Gierl [syn.: Baeomycesfungoides (Sw.) Ach.]; CO, VE, EC; 1000-3950 m Icmadophila Trevis. Ref.: López-Figueiras, 1986; Sipman, 1989. J. aversa (Nyl.) Rambold & Hertel (syn.: G!ossodium aversum Nyl.); CO, VE; 2750-4040 m
LECANORACEAE Carbonea.(Hertel) Hertel Ref.: Hertel, 1971. C. vorticosa (Fliirke) Hertel; VE; 3900-4150 m Lecanora Ach. Ref.: Hertel, 1971. L. caesiorubella Ach. subsp. glaucomodes (Nyl.) Imshang & Brodo; CO; 3200 m [Note: leg. Sipman & Aguirre 27460 (B)] L. intricata (Schrader) Ach.; CO; 3200 m [Note: leg. Sipman & Aguirre 27439 (B)] L. polytropa (Ehrh.) Rabenh.; CO, VE; 3200-4750 m
LECIDEACEAE Ref.: Hertel, 1971. Lecidea Ach. L. confluentula Müll.Arg.; VE; 4700 m L. /apicida (Ach.) Ach.; VE; 3900-4150 m L. limosa Ach.; VE; 4000-4700 m L. secernens H.Magn.; VE; 4700 m L. tesselata Fliirke; VE; 3900-415Ó m
LOBARIACEAE Lobaria (Schreb.) Hoffm. Ref.: López-Figueiras, 1986; Sipman, 1989; Yoshimura, 1984; Yoshimura & Arvidsson, 1994. L. botryoides Yoshim. & Arv.; EC; 3350-3950 m L. eren u/ata (Hook.) Trevis.; CR, PA, CO, EC; 1100-3700 m L. exornata (Zahlbr.) Yoshim. var. cora/lophora Yoshim.; CO, EC; 2000-3500 m L. pallida (Hook.) Trevis.; CR, CO, VE, EC; 1600-4000 m L. subdissecta (Nyl.) Vain.; CR, CO; 1800-3500 m L. subexornata Yoshim.; CR, CO, EC; 1100-3800 m Pseudocyphellaria Vain. Ref.: Galloway & Arvidsson, 1990; Sipman, 1989. P. aurata (Ach.) Vain.; EC; 700-3500 m P. bartlettii D.J.Galloway; EC; 3350-3600 m P. crocata (L.) Vain.; CO, EC; 1500-3750 m P. encoensis R.Sant.; EC; 3500 m P. intricata (Delise) Vain.; EC; 3400 m Sticta (Schreb.) Ach. Ref.: Gyelnik, 1938; Kalb, 1988; López-Figueiras, 1986; Nylander, 1863a, 1863b; Sipman, 1989. S. ambavillaria (Bory) Ach.; VE; 3450 m S. andensis (Nyl.) Trevis.; CO, VE; 1800-4400 m S. camelia Ach.; EC; 4000 m S.fit!iginosa (Dicks.) Ach.; CR, CO, VE, EC; 1600-4750 m S. granatensis Nyl.; CO; 2125-3750 m S. gya/ocarpa (Nyl.) Trevis.; CR, CO, VE; 2300-4325 m S. humboldtii Hook.; CO, VE; 2300-3975 m S. !imbata (Sm.) Ach.; CO; 3200 m [Note: leg. Sipman & Aguirre 27463 (B)] S. neolinita Gyeln.; CO; 3540-3800 m S. neopulmonaria Gyeln.; CO; 3200 m S. peruviana Ach.; CR, VE, EC; 3300-3350 m
46
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
S. tomentosa (Sw.) Ach.; CO, VE, EC; 1000-4330 m; in-
eludes var. dilatata (Nyl.) Hue and var. tomentosa S. weigelii Ach.; CO, VE, EC; 1250-4800 m
MEGALOSPORACEAE Megalospora Meyen Ref.: Kalb, 1990; Sipman, 1986c, 1989. M. admixta (Nyl.) Sipman; CO, EC; 2400-3975 m M. foersteri Kalb; VE; 4200 m M. tuberculosa (Fée) Sipman; CO, EC; 240-3750 m
P ANNARIACEAE Erioderma Fée Ref.: López-Figueiras, 1986. E. chilense Mont.; CR, VE; 2500-3500 m E. sorediatum D.J.Galloway & P.M.Jorg.; CR, CO, VE; 2450-3200 m [Note: CO voucher leg. Sipman & Aguirre 27392 (B)] E. verruculosum Vain.; CR, VE; 3300-3350 m [Note: CR voucher leg. Sipman 20957 (Lichenotheca Latinoamericana 21) (B)] E. wrightii Tuck.; VE; 3000-3500 m Pannaria Delise ex Bory Ref: López-Figueiras, 1986; Sipman, 1989. P. conoplea (Ach.) Bory; CO; 3100-3500 m P. rubiginosa (Ach.) Bory; CO, VE; 2500-4325 m Parmelie/la Müll.Arg. Ref.: López-Figueiras, 1986; Sipman, 1989. P. mariana (Fr.) P.M.Jorg. & D.J.Galloway [syn.: Pannaria mariana (Fr.) Mü11.Arg.]; CO; 2000-3900 m P. nigrocincta (Mont.) Müll.Arg.; CR, CO, VE; 200-4075 m Psoroma Michaux Ref.: Arvidsson, 1991; López-Figueiras, 1986; Sipman, 1989. P. hypnorum (Vahl) Gray; CO, EC; 3700-4550. P. paleaceum (Fr.) Nyl.; CO; 4150-4175 m
[VOL. 84
A. dictyorhiza (Massee) Yoshim.; VE; 2750-3600 m A. leucobates (Nyl.) Müll.Arg.; CO, VE; 2000-3600 m A. lopezii Yoshim.; unspecified; 3000-3600 m A. masonii Yoshim.; CO, VE; 2400-3750 m A. parasítica (Fée) Zahlbr.; CO, VE; 2000-3500 m Bulbothrix Hale Ref.: Hale, 1976. B. imshaugii (Hale) Hale; VE; 2600-3200 m B. laevigatula (Nyl.) Hale; CO; 2700-3500 m Canoparmelia Elix & Hale C. texana (Tuck.) Elix & Hale [syn.: Pseudoparmelia texana (Tuck.) Hale]; VE; 800-3500 m Céraria Ach. Ref.: Kamefelt, 1979; Kamefelt et al., 1993; Sipman, 1992b. C. arenaría Kamefelt; CO; 2700-3815 m C. islandica (L.) Ach.; CO; 3590-4475; includes subsp. crispiformis (Rasanen) Kamefelt and subsp. islandica C. muricata (Ach.) Eckf. [syn.: Coelocaulon muricatum (Ach.) Laundon]; VE, EC; 3500 m C. nigricans N y!.; CO; 4320 m Cetrariastrum Sipman Ref.: Sipman, 1980. C. andense Kamefelt; VE, EC; 3300-4000 m C. dubitans Sipman; CO, EC; 3300-4475 m C. ecuadoriense (R.Sant.) Sipman; CO, EC; 4250-4325 m Cetrariella Kamefelt & A.Thell Ref.: Kamefelt el al., 1993. C. delisei (Bory ex Schaer.) Kamefelt & A.Thell [syn.: Cetraria delisei (Bory ex Schaer.) Nyl.]; EC;? m
Alectoria Ach. Ref.: Marcano et al., 1994. A. imshaugii Brodo & D.Hawksw. var. venezuelensis V.Marcano & AMorales; VE; 3500-3800 m A. ochroleuca (Hoffm.) A.Massal.; CO, VE, EC; 3350-4510 m; includes var. ochroleuca and var. tropica V.Marcano & A.Morales [Note: Marcano et al. (1994) suggested that all neotropical material of Alectoria ochroleuca belongs lo var. tropica.]
Everniastrum 1:-íale ex Sipman Ref.: Kalb, 1988; Sipman, 1980, 1986a. E. arvidssonii Sipman; EC; 3000-3200 m E. catawbiense (De gel.) Hale ex Sipman; CR, CO, VE, EC; 3000-4020 m E. cirrhatum (Fr.) Hale ex Sipman; CR, CO, VE, EC; 20004150 m E. columbiense (Zahlbr.) Hale ex Sipman; CO, VE, EC; 2300-3600 m E. fragile Sipman; CR, CO, VE, EC; 3000-4400 m [Note: CR voucher leg. Sipman (B).] E. lipidiferum (Hale & M.Wirth) Hale ex Sipman; CO, VE; 3000-3750 m E. nigrociliatum (de Lesd.) Hale ex Sipman; VE; 3500 m E. planum Sipman; CR, CO, EC; 3300-3350 m E. sorocheilum (Vain.) Hale ex Sipman; CR, CO, VE, EC; 2400-4540 m E. vexans (Zahlbr. ex W.L.Cu1b. & C.F.Culb.) Hale ex Sipman; CR, CO, VE, EC; 1400-4050 m
Anzia Stizenb. Ref.: Yoshimura, 1995.
Everniopsis N y!. E. trulla (Ach.) Nyl.; EC; 3500 m
P ARMELIACEAE Re f.: Arvidsson, 1991; López-Figueiras, 1986; Sipman, 1989.
1999]
PARAMOS
Flavoparmelia Hale F. ecuadoriensis Nash, Elix & J.Johnst.; CO, EC; 3200 m [Note: For EC voucher see Arvidsson, 1991.] F. gerlachei (Zahlbr.) Hale [syn.: Pseudoparmelia gerlachei (Zahlbr.) Hale]; CO, VE; 1800-3700 m F. soredians (Nyl.) Hale [syn.: Pseudoparmelia soredians (Nyl.) Hale]; VE; 2700-3800 m
H. H. H. H. H. H. H.
Flavopunctelia Hale F.jlaventior (Stirt.) Hale; VE, EC; 2000-3400 m
H. H. H. H.
Hypogymnia (Nyl.) N y!. Ref.: Sipman & Aguirre, 1982. H. bitteri (Lynge) Ahti; CO, VE, EC; 3100-4550 m H. vittata (Ach.) Gas.; VE; 3500 m [Note: leg. Sipman 11344 (B)] Hypotrachyna (Vain.) Hale Ref.: Aguirre, 1982; Hale, 1972, 1975. H. andensis Hale; CR, CO, VE, EC; 1600-4000 m H. bogotensis (Vain.) Hale; CR, CO, VE, EC; 2200-4000 m H. brevirhiza (Kurok.) Hale; CO, VE, EC; 1400-4325 m H. caraccensis (Taylor) Hale; CR, CO, VE, EC; 1400-4300 m H. cendensis Hale & López-Fig.; VE; 2800-3400 m H. chicitae (Hale) Hale; VE; 2700-4200 m H. costaricensis (Nyl.) Hale; CO, VE, EC; 1000-3750 m H. croceopustulata (Kurok. & Hale) Hale; VE; 1400-3750 m H. degelii (Hale) Hale; CO, VE; 1400-3500 m H. densirhizinata (Kurok.) Hale; CR, CO, VE, EC; 24003900 m H. dentella (Hale & Kurok.) Hale; VE; 1600-3800 m H. ducalis (Jatta) Hale; CR, CO, VE; 1600-3450 m H. enderythraea (Zahlbr.) Hale; CO, VE; 1900-4000 m H. endoch/ora (Leight.) Hale; CO, VE, EC; 1900-3400 m H. ensifolia (Kurok.) Hale; CR, CO, VE, EC; 2800-4100 m H. erythrodes (Zahlbr.) Hale; CO, VE; 1400-3750 m H. exsplendens (Hale) Hale; CO, VE, EC; 1400-4750 m H. flavida (Zahlbr.) Hale; CO, VE; 1200-3750 m H. gigas (Kurok.) Hale; CR, CO, VE, EC; 2200-3900 m H. gondy/ophora (Hale) Hale; VE; 1800-3600 m H. imbricatula (Zahlbr.) Hale; CR, CO, VE; 1100-3730 m H. immaculata (Kurok.) Hale; CO, VE; 1800-3780 m H. laevigata (Sm.) Hale; CR, CO, VE, EC; 1300-3935 m H. lineariloba (Kurok.) Hale; VE; 2400-3500 m H. livida (Taylor) Hale; VE; 2100-3450 m H. lopezii Hale; CR, CO, VE; 2800-4050 m H. microblasta (Vain.) Hale; CR, CO, VE, EC; 1400-3975 m H. monilifera (Kurok.) Hale; CO, VE; 2350-3600 m H. neodissecta (Hale) Hale; VE; 1300-3300 m H. neojlavida Hale; VE; 3200 m H. obscurella (Vain.) Hale; CO, VE; 1900-3500 m H. oostingii (Dey) Hale; VE; 3300-3500 m H. osteoleuca (Nyl.) Hale; CO, VE, 1500-3500 m H. paramensis W.L.Culb. & C.F.Culb.; VE; 2400-3750 m H. partita Hale; CR, CO, VE; 2400-3750 m H. physcioides (Nyl.) Hale; CR, CO, VE, EC; 1900-4100 m H. physodalica (Hale) Hale; CO, VE, EC; 2950-4050 m H. primitiva Hale & López-Fig.; CO, VE; 3100-4000 m H. producto Hale; CO, VE, EC; 1300-3500 m
47
prolongata (Kurok.) Hale; CR, CO, VE; 2300-3800 m protenta Hale; VE; 1300-4000 m protoboliviana (Hale) Hale; CR; 3300 m pseudosinuosa (Asahina) Hale; VE; 1500-3750 m pulvinata (Fée) Hale; CR, CO, VE, EC; 1200-4300 m reducens (Nyl.) Hale; CR, CO, VE; 2600-4175 m revoluta (Fliirke) Hale; CO; 1700-3400 m [Note: leg. Aguirre & Sipman 5322 (B)] rockii (Zahlbr.) Hale; CO, VE, EC; 1500-4050 m sinuosa (Sm.) Hale; CR, CO, VE, EC; 1200-4540 m sublaevigata (Nyl.) Hale; CO, EC; 200-3800 m subsaxatilis (de Lesd.) Hale; VE; 3100-4050 m
Melanelia Essl. M. panniformis (Nyl.) Essl.; VE; 4200 m Oropogon Th.Fr. Ref.: Esslinger, 1989. O. aliphaticus Essl.; PA, VE; 2500-3300 m O. americanus Essl.; CR, CO, VE, EC; 3300-4100 m O. atranorinus Essl.; CR, VE; 2000-3490 m O. barbaticus Essl.; CR, CO, VE, EC; 2880-4200 m O. bicolor Essl.; CR, PA, CO, VE, EC; 2600-4300 m O. byssaceus Essl.; VE; 4050 m O. colibor Essl.; CR, PA, VE; 2400-4000 m O. diffractaicus Essl.; CR, PA, VE; 1600-4000 m O.fissuratus Essl.; CR, VE; 1600-3600 m O.formosanus Asahina; PA, CO, VE, EC; 2100-4200 m O. granulosus Essl.; VE, EC; 2500-3500 m O. halei Essl.; CR, VE; 3000-3500 m O. lopezii Essl.; VE; 2250--3300 m [Note: leg. Sipman 11222 (B)] O. lorobic Essl.; CR, PA, VE; 2450-3600 m O. loxensis (Fée) Th.Fr. (sensu Essl.); CR, PA, CO, VE, EC, PE; 2400-4200 m O. macilentus Essl.; EC; 3450 m O. maurus Essl.; VE; 3200-3600 m O. parietinus Essl.; VE; 3500 m O. pendulus Essl.; EC; 4300-4450 m O. sperlingii Essl.; CR, CO, VE, EC; 2400-4100 m O. striatulus Essl.; VE; 2400-3500 m Parmelina Hale P. versiformis (Kremp.) Hale; VE; 2600-3450 m Parmelinopsis Elix & Hale P. cleefii (Sipman) V.Marcano & Sipman; CO, VE; 34504300 m P. cryptochlora (Vain.) Elix & Hale [syn.: Parmelina cryptochlora (Vain.) Hale]; VE; 1300-3400 m P. swinscowii (Hale) Elix & Hale (syn.: Parmelina swinscowii Hale); CO, VE, EC; 3200-4550 m Parmotrema A.Massal. Ref.: Aguirre, 1982; Hale, 1965. P. arnoldii (DuRietz) Hale; CO, VE, EC; 1950-3935 m P. bangii (Vain.) Hale; EC;? m [Note: leg. Harris (NY)] P. chinense (Osbeck) Hale & Ah ti; EC;? m [Note: leg. Harris (NY)]
48
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
conferendum Hale; VE; 1200-3300 m crinitum (Ach.) M.Choisy; CO, VE, EC; 1400-3600 m eborinum (Hale) Hale; VE; 200-3450 m ecrinitum Hale; VE; 2200-3450 m fasciculatum (Vain.) Hale (syn.: Parmelia fasciculata Vain.); CO, VE; 3300-3780 m P.fractum (Hale) Hale; CO, VE, EC; 2300-3950 m P. mellissii (Dodge) Hale; VE; 1225-3450 m P. rampoddense (Nyl.) Hale; VE; 1200-3300 m P. robustum (Degel.) Hale (syn.: Parmotrema dilatatum auct.); CO, VE; 1500-3800 m P. virescens Hale; VE; 1400-3500 m P. P. P. P. P.
Protoparmelia M.Choisy Re f.: Hertel, 1971. P. badia (Pers.) Hafellner; VE; 3900-4740 m P. picea (Dicks.) Nyl.; CO; 3200 m [Note: leg. Sipman & Aguirre 27440 (B)] Psiloparmelia Hale P. distincta (Nyl.) Hale [syn.: Xanthoparmelia distincta (Nyl.) Hale]; EC; 4100 m Punctelia Krog P. rudecta (Ach.) Krog; VE, EC; 1200-3200 m P. stictica (Duby) Nyl.; CR, CO, VE, EC; 2400-4080 m P. subrudecta (Nyl.) Krog; VE; 900-3400 m Rimelia Hale & A.Fletcher R. cetrata (Ach.) Hale & A.Fletcher [syn.: Parmotrema cetratum (Ach.) Hale]; CO, VE; 400--4200 m R. reticulata (Taylor) Hale & A.Fletcher [syn.: Parmotrema reticulatum (Taylor) Hale]; CO, VE, EC; 1000-3650 m Tuckneraria Randlane & A.Thell Ref.: Kalb, 1990. T. laureri (Kremp.) Randlane & A.Thell (syn.: Cetraria laureri Kremp.); CO, VE; 3500--4200 m Usnea Dill. ex Adans. Ref.: Kalb, 1988; Lindan, 1912; Motyka, 1936-1938; Vainio, 1899a; Walker, 1985. U. acanthella (Lamb) F.J.Walker; VE, EC; 3450--4500 m U. angulata Ach.; CO; 2000-3500 m U. bogotensis Vain.; CO, VE, EC; 2200-3950 m U. durietzii Motyka; PA, CO, VE, EC; 2800--4750 m U. jelskii Motyka; CO; 3500-4000 m U. moreliana Motyka; CO; 2740-3800 m U. sphacelata R.Br.; CO, VE, EC; 4000-4900 m U. spinul!fera (Vain.) Motyka; CO; 3600-4250 m Xanthoparmelia (Vain.) Hale Ref.: Nash et al., 1995. X arvidssonii Nash & Elix; EC; 3200-3900 m X conspersa (Ach.) Hale; VE; 3500 m X cotopaxiensis Nash, Elix & J.Johnst.; EC; 3400 m X cumberlandia (Gyeln.) Hale; VE; 2800--4400 m X lopezii Nash & Elix; VE; 4100 m
X X X X X X X X X X X X X
[VOL. 84
microspora (M체li.Arg.) Hale; VE, EC; 2100--4200 m mougeotii (Schaer.) Hale; CO, VE, EC; 2900--4200 m neowyomingica Hale; EC; 2600-3900 m sipmanii Nash & Elix; VE; 4100 m somloensis (Gyeln.) Hale; VE; 3800 m standaertii (Gyeln.) Hale; VE; 3700 m subsorediata Hale; VE, EC; 3000-4200 m subulcerosa Nash & Elix; EC; 2300-3840 m succedans Elix & J.Johnst.; EC; 4200 m tasmanica (Hook.f. & Taylor) Hale; VE; 3200 m ulcerosa (Zahlbr.) Hale; VE; 2100-3700 m vagans (N y l.) Hale; EC; 3200 m wildeae (Dodge) Hale; EC; 1680-3300 m
PELTIGERACEAE Nephroma Ach. Ref.: L처pez-Figueifas, 1986; Sipman, 1989, 1990. N. helveticum Ach.; CR, CO, VE, EC; 2700-3900 m Peltigera Willd. Ref.: Arvidsson, 1991; L처pez-Figueiras, 1986; Sipman, 1989; Vitikainen, 1994. P. austroamericana Zahlbr.; CO, VE, EC; 1550--4000 m P. canina (L.) Willd.; VE; 2400-3800 m P. didactyla (With.) J.R.Laundon [syn.: P. spuria (Ach.) DC.]; CO; 1900--4680 m P. dolichorhiza (Nyl.) Nyl.; CO; 1000-3750 m P. lacinia/a (Merril ex Riddle) Gyeln.; CO, VE, EC; 15004200 m P. pulverulenta (Taylor) Nyl.; CR, CO, VE, EC; 2300-4400 m P. rufescens (Weiss) Humb.; VE; 1750--4050 m P. soredians Vitik.; CO; 3200-3750 m P. ulcera/a M체ll.Arg.; CO, VE; 1700--4150 m P. vainioi Gyeln.; CO, VE; 2000-3300 m So/orina Ach. Re f.: Arvidsson, 1991. S. saccata (L.) Ach.; EC; ? m S. spongiosa (S m.) Anzi; CO; 3675 m [Note: leg. Cleef5432 (B)]
PERTUSARJ!ACEAE Ref.: Sipman, 1986b, 1989. Pertusaria DC. P. acroscyphoides Sipman; CO; 3830--4450 m P. culbersonii Vezda; CO, VE; 3500--4450 m
PHL YCTIDACEAE Phlyctis Wallr. Ref.: Sipman, 1989. P. endecamera (Nyl.) Nyl. [syn.: Phlyctella endecamera (Nyl.) Nyl.]; CO; 2400-3500 m
PHYSCIACEAE Buellia De Not. Ref.: Hertel, 1971.
1999]
PARAMOS
B. nivalis (Bagl. & Carestía) Hertel ex Hafellner [syn.: Polyschistes nivalis (Bagl. & Carestía) Keissl.]; VE; 4250 m Dirinaria (Tuck.) Clem. Ref.: López-Figueiras, 1986. D. applanata (Fée) Awas.; VE; 30-2200(-3452) m
49
PILOCARP ACEAE Byssoloma Trevis. B. subdiscordans (Nyl.) P.James; CO; 300-3200 m [Note: leg. Aguirre & Sipman 5145 (B)]
PORPIDIACEAE Hafellia Kalb, H.Mayrhofer & Scheid. Re f.: López-Figueiras, 1986. H. dissa (Stirt.) H.Mayrhofer & Sheard [syn.: Buellia callispora (Knight) Stein]; VE; 3300 m
Ref.: Hertel, 1971. Porpidia Korb. P. crustulata (Ach.) Herte! & Knoph; VE; 3500 m
PSORACEAE Heterodermia Trevis. Ref.: Aptroot, 1989; López-Figueiras, 1986. H. albicans (Pers.) Swinscow & Krog; EC; ? m [Note: leg. Sipman 7777 (B)] H. antillarum (Vain.) Swinscow & Krog; VE; 1400-3400 m H. casare/liana (A.Mass.) Trevis.; CR, CO, VE; 1000--3750 m H. circinalis (Zahlbr.) W.A. Weber; CR, CO, VE, EC; 23003800 m H. corallophora (Tayl.) Skorepa; VE; 600-3300 m H. ga/actophylla (Tuck.) Trevis.; CO; 2120-3730 m H. hypoleuca (Mühl.) Trevis.; CO; 2200-3200 m H. isidiophora (Vain.) Awas.; VE, EC; 1200-3450 m H.japonica (Sato) Swinscow & Krog; VE; 900-3500 m H. /eucome/os (L.) Poelt [syn.: H. boryi (Fée) Kr.P.Singh & S.R.Singh]; CO, VE, EC; 400-3700 m H. lutescens (Kurok.) Follmann & Redon; CR, CO; 16003750 m H. magellanica (Zahlbr.) Swinscow & Krog; VE, EC; 13003750m H. obscura/a (Nyl.) Trevis.; VE; 700-3900 m H. podocarpa (Bél.) Awas.; VE; 1300-3500 m H. speciosa (Wulf.) Trevis.; CO, VE; 300-3730 m H. verrucifera (Kurok.) W.A.Weber; VE; 1100-3450 m . H. vulgaris (Vain.) Follmann & Redon; EC;? m [Note: leg. Harris (NY)] Hype1physcia Müll.Arg. H. adglutinata (F!Orke) Mayrhofer & Poelt; EC;? m [Note: leg. Sipman 7785 (B)]
Ref.: Hertel, 1971; Kalb, 1988. Psora Hoffm. P. icterica (Mont.) Müll.Arg.; VE, EC; 4100-4500 m
RAMALINACEAE Ref.: López-Figueiras, 1986; Kalb, 1988; Marcano & Morales Méndez, 1993. Ramalina Ach. R. asahinae W.Culb. & C.Culb.; VE; 600-3500 m R. camptmpora Nyl.; VE; 3200-3600 m R. ce/astri (Spreng.) Krog & Swinscow; VE; 1200-3500 m R. cochlearis Zahlbr.; VE; 1700-3800 m R. complanata (Sw.) Ach.; VE; 1100-3500 m R. cumanensis Fée; VE, EC; 2100-3550 m R. dictyota Dodge & Vareschi; VE, EC; 2600-3500 m R. puiggarii Müll.Arg.; VE; 3100-3500 m R. pusiola Müii.Arg.; VE; 900-3500 m R. rectangularis Nyl.; VE; 1100-3350 m R. tenuissima V.Marcano & A. Morales; VE; 2200-3400 m R. usnea (L.) R.H.Howe; VE; 1100-3500 m
RHIZOCARP ACEAE Ref.: Hertel, 1971. Rhizocarpon Ramond ex DC. R. /eptolepis Anzi; VE; 3500 m R. superficiale (Schaer.) Vain.; VE; 4250 m
SOLORINELLACEAE Phaeophyscia Moberg Re f.: López-Figueiras, 1986. P. endococcinodes (Poelt) Essl.; VE; 2000-4050 m P. hispidula (Ach.) Moberg; VE, EC; 1750-3450 m Physcia (Schreb.) Michaux Ref.: Moberg, 1990. P. cinerea Moberg; VE, EC; 2200-3600 m P. lopezii Moberg; VE; 2200-3600 m P. poncinsii Hue; VE; 200-3600 m Pyxine Fr. P. limbulatd Müli.Arg.; VE; 3800-4200 m
Ref.: Vézda & Poelt, 1991. Gyalidea Lettau G. costaricensis Vézda & Hafellner; CO; 3300-3390 m G. hya/inqcens (Nyl.) Vézda var. hyalinescens; EC; 20003550 m
SPHAEROPHORACEAE Ref.: Arvidsson, 1991; López-Figueiras, 1986; Sipman, 1990. Bunodophoron A.Massal. B. me/anocarpum (Sw.) Wedin [syn.: Sphaeroplzorus melanocarpus (Sw.) DC.]; CO, VE, EC; 1800-4750 m
50
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN ·
STEREOCAULACEAE Ref.: Arvidsson, 1991; Boekhout, 1982; López-Figueiras, 1986; Sipman, 1986b, 1989.
Stereocaulon Hoffm. S. atlanticum (Lamb) Lamb; CR, CO, VE; 2700-4500 m S. claviceps Th.Fr.; CO; 3750-4300 m S. crambidiocephalum Lamb; CR, CO, VE, EC; 3100-4340 m S. delisei Bory ex Duby; CR, CO; 3350-4340 m S. didymicum Lamb; VE; 2000-3400 m [Note: 1eg. Sipman 11212 (U)]
S. glareosum (Savicz) H.Magn.; CO, EC; 4000-4530 m S. globisorum Sipman; CO, EC; 3750-4340 m S. meyeri Stein; CO, VE, EC; 2450-4300 m S. microcarpum Müll.Arg.; VE, EC; 1400-4050 m S. novogranatense Lamb; CO, EC; 1700-3750 m . S. obesum Th.Fr.; CO, VE, EC; 2450-4160 m S. pachycephalum Vain.; CO; 2500-4200 m S. pityrizans N y l.; CO, VE, EC; 2300-3500 m . S. pomiferum P.A.Duvign.; CO, VE, EC; 1500-3750. in S. ramulosum Raeusch.; CR, CO, VE, EC; 1800-4500 m S. strictum Th.Fr.; CR, CO, VE, EC; 1700-3900(-4500) m; inc1udes var. compressum (Nyl.) Lamb ex Vezda. and var. strictum S. tomentosum Th.Fr. [syn.: S. myriocarpum Th.Fr.]; CR, CO, VE, EC; 2300-4400 m S. verruciferum Nyl. var. surreptans (Lamb) Lamb; EC; 3400-4500 m S. vesuvianum Pers.; CR, CO, VE, EC; 2100-4750 m
TELOSCHISTACEAE Caloplaca Th.Fr. C. brebissonii (Fée) R.Sant. ex Hafellner & Poe1t; CO, EC; 2100-3400 m [Note: CO voucher 1eg. Aguirre & Sipman 5818 (B)] C. calicioides P.M.Jorg.; VE; 3500 m Te/oschistes Norman Ref.: López-Figueiras, 1986. T. ch1ysophthalmus (L.) Th.Fr.; VE, EC; 2900-3450 m T. exilis (Michx.) Vain.; EC;? m [Note: leg. Harris (NY)] T.jlavicans (Sw.) Norman; VE, EC; 700-3500 m T. hypoglaucus (Nyl.) Zah1br.; EC; ? m [Note: 1eg. Sipman 7792 (B)] Xanthoria (Fr.) Th.Fr. Ref.: López-Figueiras, 1986. X elegans (Link) Th.Fr.; VE; 3900-4250 m
THELOTREMATACEAE Diploschistes Norman Re f.: Arvidsson, 1991 ; López-Figueiras, 1986; Sipman, 1989. D. cinereocaesius (Sw. ex Ach.) Vain.; CR CO VE EC· 2400-4450 m ' ' ' ' D. diacapsis (Ach.) Lumbsch; CR; 3400 m [Note: 1eg. Sipman & Chaverri 20705 (B)] D. gypsaceus (Ach.) Zah1br.; VE; 3450-3500 m
[VOL. 84
D. muscorum (Scop.) R.Sant. subsp. bartlettii Lumbsch; CO; 1900-4300 m The/otrema Ach. Ref.: Sipman, 1989. T. lepadinum (Ach.) Ach.; CO; 2700-3730 m
TRAPELIACEAE Placopsis (Nyl.) Linds. Ref.: Ka1b, 1988; López-Figueiras, 1986. P. gelida (L.) Lindsay; CO, VE, EC; 2100-3700 m P. parellina (Nyl.) Lamb; EC; 3400 m Placynthiella E1enkin Ref.: Sipman, 1989. P. icmalea (Ach.) Coppins & P.James [syn.: Saccomorpha icmalea (Ach.) Clauzade & Cl.Roux]; CO; 4350 m Trapelia M.Choisy Ref.: Hertel, 1971. T. coarctata (Tum.) M.Choisy; VE, EC; 1600-3900 m T. mooreana (Caroll) P.James [syn.: Trapelia torellii (Anzi) Herte1]; CO, VE; 2000-3500 m T. obtegens (Th.Fr.) Herte1; VE; 3500 m Trapeliopsis Herte1 & Gotth.Schneid. Ref.: Hertel, 1971, 1973; Sipman, 1989. T. glauco/epida (Nyl.) Gotth.Schneid.; CR, CO; 3130-4400 m T. subconcolor (Anzi) Herte1; CO, VE; 3500-4000 m
UMBILICARIACEAE Ref.: López-Figueiras, 1986; Marcano, 1994; Sipman, 1989, 1992a.
Umbilicaria Hoffm. U. africana (Jatta) Krog & Swinscow; CO, VE, EC; 30004750m U. calvescens Nyl.; VE, EC; 3000-4100 m U. hap/ocarpa Nyl.; EC; ? m [Note: herb. Aptroot] U. leprosa (Zah1br.) Frey; CR, CO, EC; 3200-4540 m U. mifafensis V.Marcano & A.Morales; VE; 4115 m U. nylanderiana (Zah1br.) H.Magn.; VE, CO, EC; 38004750m U. po/yphylla (L.) Baumg.; CO, VE; 3200-4330 m U. po/yrrhiza Ach.; CO, VE; 2700-3850 m U. subca/vescens Sipman; CO, VE; 2450-4350 m
VERRUCARIACEAE Catapyrenium F1ot. Ref.: Breuss, 1993. C. exaratum Breuss; EC; 4500 m C. lachneoides Breuss; EC; 3300-4500 m C. squamulosum (Ach.) Breuss var. squamulosum; EC; 3900-4500 m Normandina Nyl. Ref.: López-Figueiras, 1986; Sipman, 1989. N. pulchella (Borr.) Nyl.; CO, VE, EC; 1750-3975 m
1999]
PARAMOS
INCERTAE SEDIS Cystacaleus Thwaites Ref.: Sipman, 1990. C. ebeneus (Dillwyn) Thwaites; CO, VE; 3000-4500 m Lepraria Ach. Ref.: Sipman, 1989. L. caesiaatra (de Lesd.) J.R.Laundon (syn.: L. neglecta Auct.); CO; 4300-4500 m Lepracaulan Lamy Ref.: Arvidsson, 1991; Lamb & Ward, 1974; LópezFigueiras, 1986; Marcano etal., 1997; Sipman, 1989. L. albicans (Th.Fr.) Nyl. ex Hue; CO, EC; 2800-4475 m L. cangestum (Nyl.) Lamb & A.Ward; CO, VE, EC; 28004400 m L. gracilescens (Nyl.) Lamb & A.Ward; VE, EC; 3500-4450 m L. micrascapicum (Vill.) Gams ex D.Hawksw.; VE; 35003600m Lepra/ama Crombie Ref.: Leuckert & Kümmerling, 1991. L. vauauxii (Hue) Laundon; CO, EC; 4300 m Racadium Pers. R. rupestre Pers.; VE; 3000-4500 m Siphula Fr. Ref.: Arvidsson, 199l;.López-Figueiras, 1986; Sipman, 1984. S. ceratites (Wahlenberg) Fr.; CO; 3100 m [Note: leg. Aguirre & Sipman 5857 (B)] S.fastigiata (Nyl.) Nyl.; CR, CO, VE, EC; 3100-4560 m S. pterulaides N y l.; CR, CO, VE, EC; 2400-4560 m Thamnalia Ach. ex Schaerer Ref.: Arvidsson, 1991; López-Figueiras, 1986; Sipman, 1989. T. vermicularis (Sw.) Schaer. emend. Asahina; CR, CO, VE, EC; 2800-4550 m
Lichenized Basidiomycetes MERULIACEAE Dictyanema Agardh ex Kunth Ref.: López-Figueiras, 1986; Sipman, 1989. D. glabratum (Spreng.) D.Hawksw. [syn.: Cara pavania (Sw.) Fr.]; CO, VE, EC; 240-4550 m D. sericeum (Sw.) Berkh.; VE; 1900-3300 m
TRICHOLOMATACEAE Omphalina Qué!. Ref.: Arvidsson, 1991; Jorgensen, 1989.
51
O. ericetarum (Pers. ex Fr.) M.Lange; VE; 3500 m O.faliacea P.M.Jorg.; CO, VE, EC; 3555-4250 m Phytacanis Bory Ref.: Redhead & Kuyper, 1987. P. aurantiaca (Redhead & Kuyper) Redhead & Kuyper (syn.: Batrydina aurantiaca Redhead & Kuyper); CO; 3000-4000 m P. !abata (Redhead & Kuyper) Redhead & Kuyper (syn.: Batrydina !abata Redhead & Kuyper); CO; 3000-4000 m
Acknowledgments The kind cooperation ofR. C. Harris (NY) andA. Aptroot (Baarn), who provided unpublished inforrnation oftheir Ecuadorean collections, is gratefully acknowledged.
Literature Cited in Checklist of Lichens Aguirre C., J. 1982. Contribución al conocimiento de los líquenes de Colombia-U. Observaciones acerca de Hypatrachyna y Parmatrema (Líquenes: Parmeliaceae). Bol. Dept. Biol., Fac. de Cienc. 1(4): 29-44. Ahti, T. 1961. Taxonomic studies on reindeer lichens (Cladania, subgenus Cladina). Ann. Soc. Zool. Bot. Fenn. Vanamo 32(1): 1-160, pi. 1-44. --~. 1986. New species of reindeer lichens (Cladina). Ann. Bot. Fenn. 23: 221-227. --~ & M. P. Marcelli. 1995. Taxonomy ofthe Cladania verticillaris complex in South America. Biblioth. Lichenol. 58: 5-26. --~ & S. Stenroos. 1986. A revision of Cladania sect. Cacciferae in the Venezuelan Andes. Ann. Bot. Fenn. 23: 229-238. Aptroot, A. 1989. Studies on Colombian cryptogams XL. The family Pyxinaceae (lichenized fungi). Proc. Kon. Ned. Akad. Wetensch., Ser. C, 92: 269-280. Arvidsson, L. 1982. A monograph of the lichen genus Caccacarpia. Opera Bot. 67: 1-96, figs. 1-63. --~. 1991. Lichenological studies in Ecuador. Pp. 123134. In: D. J. Galloway (ed.), Tropical lichens: Their systematics, conservation, and ecology. Systematics Association Special Volume 43. Clarendon Press, Oxford. Boekhout, T. 1982. Studies on Colombian cryptogams XVIII. The genus Stereacaulan (Schreber) Hoffmann (Lichenes). J. Hattori Bot. Lab. 53: 438-511. Brako, L. 1991. Phyllapsara (Bacidiaceae). Fl. Neotrop. Monogr. 55: 1-67. Breuss, O. 1993. Catapyrenium (Verrucariaceae) species from South America. PI. Syst. Evol. 185: 17-33. Degelius, G. 1974. The lichen genus Ca!iema with special reference to the extra-European species. Symb. Bot. Upsal. 20(2): 1-215. Esslinger, T. L. 1989. Systematics of Orapagan (Alectoriaceae) in the New World. Syst. Bot. Monogr. 28: 1-111. Filson, R. B. 1981. A revision ofthe lichen genus Cladia. Unpubl. list of specimens.
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MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
Galloway, D. J. & L. Arvidsson. 1990. Studies in Pseudocyphellaria (Lichenes) II. Ecuadorian species. Lichenologist 22: 103-135. Gierl, C. & K. Kalb. 1993. Die Flechtengattung Dibaeis. Eine Übersicht über die rosafrüchtigen Arten von Baeomyces sens. la t. nebst Anmerkungen zu Phyllobaeis gen. nov. Herzogia 9: 593-645. Gyelnik, V. (Kiifaragó-). 1938. Fragmenta lichenologica I. Lilloa 3: 49-80. Hale, M. E. 1965. A monograph of Parmelia subgenus Amphigymnia. Contr. U.S. Natl. Herb. 36(5): 193-358, pi. 1-16, figs. 1-29. - - - . 1972. New species of Parme/ia sect. Hypotrachyna. Phytologia 22: 433-438. ---·. 1975. A revision ofthe lichen genus Hypotrachyna (Parmeliaceae) in tropical America. Smithsonian Contr. Bot. 25: i-iii, 1-73, figs. 1-20. - - - . 1976. A monograph ofthe lichen genus Bulbothrix Hale (Parmeliaceae). Smithsonian Contr. Bol. 32: 1-29, figs. 1-7. Hawksworth, D. L. 1985. A redisposition ofthe species referred to the ascomycete genus Microthelia. Buli: Brít. . Museum (Nat. Hist.), Bot. Ser. 14(2): 43.:-181. Hertel, H. 1971. Über holarktische Krustenflechten aus den venezolaníschen Anden. Willdenowia 6: 225-272. - - - . 1973. Beitrage zur Kenntnis der Flechtenfamilie Lecideaceae V. Herzogia 2: 479-515. J0rgensen, P. M. 1989. Omphalína foliacea, a new basídíolichen from America. Nordic J. Bot. 9: 89-95. Kalb, K. 1988. (Schedae ad) Lichenes Neotropici, Fascikel X (No. 401-450), p. 1-16. Neumarkt/Opf., Germany. - - - . 1990. (Schedae ad) Lichenes Neotropicí, Fascikel XI (No. 451-475), p. 1-12. Neumarkt/Opf., Germany. Kamefelt, I. 1979. The brown fruticose species ofCetraria. Opera Bot. 46: 1-150, figs. 1-74. ---,J.-E. Mattson & A. Thell. 1993. The 1ichen genera Arctocetraria, Cetraria, and Cetrariella (Parmeliaceae) and their presumed evolutionary affinities. Bryologist 96: 394-404. Lamb, I. M. & A. Ward. 1974. A preliminary conspectus of the species attributed to the imperfect Jichen genus Leprocau/on Nyl. J. Hattori Bol. Lab. 38:499-553, figs. 1-11. Leuckert, Ch. & H. Kümmerling. 1991. Chemotaxonomische Studien in der Gattung Lepro/oma Nyl. ex Crombie (Líchenes). Nova Hedwigia 52: 17-32. Lindau, G. 1912. Beitrag zur Kenntnis der Flechten von Kolumbien. Mém. Soc. Neuch. Sci. Nat. 5: 57-66. López-Figueiras, M. 1986. Censo de macrolíquenes venezolanos de los estados Falcón, Lara, Mérida, Táchira y Trujillo. Universidad de Los Andes, Facultad de Farmácia, Mérida, Venezuela. Marcano, V. 1994. Revisión del género Umbílicaria (Ascomicetes liquenizados) en Venezuela. Emstia 4: 21-35. - - - & A. Morales Méndez. 1993. Ramalina reducto Krog et Swinscow, especie de liquen de los Andes venezolanos, nueva para America. Ernstia 3: 19-26. ---,A. Morales Méndez & L. Martínez. 1994. El género Alectoria Ach. (Ascomicetes líquenizados) en los Andes venezolanos. Emstia 4: 89-100. - - - , - - - , H . Sipman & L. Calderón. 1996. A first checklist ofthe lichen-forming fungi ofthe Venezuelan Andes. Tropical Bryology 12: 193-235. - - - , L. Galiz, S. Mohali, A. Morales Méndez & E. Palacios-Prü. 1997. Revisión del género Leprocau/on
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Nyl. ex Lamy (Lichenes Imperfecti) en Venezuela. Tropical Bryology 13: 47-56. · Moberg, R. 1990. The Jichen genus Physcia in Central and South America. Nordic J. Bot. 10: 319-342. Motyka, J. 1936-1938. Lichenum genus Usnea studium monographicum. Pars systematica [Lwow] 1: 1-304 (1936); 2(1): 305-560 (1937); 2(2): 561-651, i-iv (1938). Müller Argoviensis, J. 1879. Lichenes aequinoctialiamericani, a el. Ed. André, annis 1875-1876, praesertim in editioribus Ecuador (E) et in Nova Granata (N.Gr.) lecti. RevueMycol. 1879(4): 163-171 (7-15). Nash, T. H., III, C. Gries & J. A. Elix. 1995. A revision of the lichen genus Xanthoparmelia in South America. Biblioth. Lichenologica 56: 1-158. Nylander, W. 1863a. Lichenographiae novo-granatensis prodromus. Acta Soc. Sci. Fenn. 7: 415-504 (pp. 1-90 · of separate edition). -·- - . 1863b. Lichens in J. Triana & J. E. Planchan, Prodromus florae novo-granatensis. Ann. Sci. Nat. Bol. IV, 19: 285-382; 20: 228-279 (p. 1-148 of separate edition). Redhead, S. A. & Th. W. Kuyper. 1987. Lichenized agarics: Taxonomic and nomenclatura! riddles. Pp. 319-349. In: G. A. Laursen, J. F. Ammirati & S. A. Redhead (eds.), Arctic and alpine mycology II. Environmental Science Research 34. Santesson, R. 1952. Foliicolous lichens I. A revision ofthe taxonomy ofthe obligately foliicolous, lichenized fungi. Symb. Bot. Upsal. 12(1): 1-590, pi. 1, figs. 1-92. Sipman, H. J. M. 1980. Studies on Colombian cryptogams X. The genus Everniastrum Hale and related taxa (Lichenes). Proceed. Kon. Ned. Akad. Wetensch., Ser. e, 83: 333-354. - - - . 1984. Lichens ofthe Buritaca-La Cumbre transect, Sierra Nevada de Santa Marta, Colombia. Pp. 185-188. In: T. Van der Hammen & P. M. Ruiz (eds.), La Sierra Nevada de Santa Marta (Colombia). Transecto BuritacaLa Cumbre. Studies of Tropical Andean Ecosystems 2. J. Cramer, Berlín. - - - . 1986a. Notes on the lichen genus Everniastrum (Parmeliaceae). Mycotaxon 26: 235-251. - - - . 1986b. Three new lichens from Colo,mbia. Willdenowia 16: 279-284. - - - . 1986c. Additional notes on the lichen family Megalosporaceae. Willdenowia 15: 557-564. - - - . 1987. Distribution patterns of the lichen genus Stereocau/on in the northern Andes. P. 320. In: W. Greuter, B. Zimmer & H. D. Behnke (eds.), Abstracts of the General Lectures, Symposium Papers and Posters presented at the XIV Intemational Botanical Congress, Berlín. - - - . 1989. Lichen zonation in the Parque Los Nevados transect. Pp. 461-483. In: T. Van der Hammen, S. Díaz P. & V. J. Alvarez (eds.), La Cordillera Central Colombiana. Transecto Parque Los Nevados (Segunda parte). Studies of Tropical Andean Ecosystems 3. J. Cramer, Berlín. - - - , 1990. Lichenotheca Latinoamericana a museo botánico berolinensi edita, fasciculum primum. Willdenowia 19: 543-551. - - - . 1992a. The genus Umbilicaria (lichenized ascomycetes) in Colombia. Nova Hedwigia 54: 63-75. - - - . 1992b. The origin ofthe lichen flora ofthe Colombian páramos. Pp. 95-109. In: H. Balslev & J. L. Luteyn
¡\
1
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PARAMOS
(eds.), Páramo: An Andean ecosystem under human influence. Academic Press, London. - - - & J. Aguirre C. 1982. Contribución al conocimiento de los líquenes de Colombia-l. Clave genérica para los líquenes foliosos y fruticosos de los páramos colombianos. Caldasia 13(64): 603-634. - - - & A. M. Cleef. 1979. Studies on Colombian cryptogams V. Taxonomy, distribution and ecology of macrolichens of the Colombian páramos: l. Cladonia subgenus Cladina. Proceed. Kon. Ned. Akad. Wetensch., Ser. C, 82: 223-241. Stenroos, S. 1989a. Taxonomic revision of the Cladonia minia/a group. Ann. Bot. Fenn. 26(3): 237-261. - - - . 1989b. Taxonomy ofthe Cladonia coccifera group 2. Ann. Bot. Fenn. 26(3): 307-317. Tibell, L. 1996. Caliciales. Fl. Neotrop. Monogr: 69: 1-79. Vainio, E. A. 1894. Monographia Cladoniarum universalis II (partie systématique et descriptive II). Acta Soc. Fauna Fl. Fenn. 10: 1-499. - - - . 1899a. Lichenes novi rarioresque, ser. l. Beibl. Hedwigia 38(3): (121)-(125). - - - . 1899b. Lichenes novi rarioresque, ser. II. Beibl. Hedwigia 38(3): (186)-(190).
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Vezda, A. & J. Poelt. 1991. Die F1echtengattung Gyalidea Lett. ex Vezda (Solorinellaceae), eine Übersicht mit Bestimmungsschlüssel. Nova Hedwigia 53: 99-113. Vitikainen, O. 1994. Notes on sorne Peltigera of the Neotropics. Acta Bot. Fenn. 150: 217-221. Vries, B. G. de & H. J. Sipman. 1984. Studies on Colombian cryptogams XXI. The lichen genus Baeomyces in Colombia and Venezuela. Proc. Kon. Ned. Akad. Wetensch., Ser. C, 87: 235-246. Walker, F. J. 1985. The lichen genus Usnea subgenus Neuropogon. Bull. Brit. Mus. (Nat. Hist.), Bot. 13(1): 1-130. Yoshimura, I. 1984. Taxonomic studies on Lobaria crenulata and its allies. J. Hattori Bot. Lab. 57: 57-66. - - - . 1994. Taxonomy and chemistry of the Lobaria crenulata group in Ecuador. Acta Bot. Fenn: 150: 223-233. - - - . 1995. The lichen genus Anzia (Parmeliaceae, Lecanorales) in Central and South America. Pp. 377387. In: F. J. A. Daniels, M. Schulz & J. Peine (eds.), Flechten Follmann, Contributions to lichenology in honour of Gerhard Follmann. Koeltz, Koenigstein. - - - & L. Arvidsson. 1994. Taxonomy and chemistry of the Lobaria crenulata group in Ecuador. Acta Bot. Fenn. 150:223-233.
Mosses Steven P. Churchill and Dana Griffin III
Mosses are a conspicuous element of two neotropicallife zones: montan e forests, particular!y cloud forests, and páramo. This contribution focuses on the taxonomic diversity ofmosses found in the pár(lmo. There are approximately 51 families, 163 genera, and 543 species presently recorded in our list ofpáramo mosses. The 1Olargest families and genera of páramo mosses account for 66% and 34%, respective! y (Table III). Endemism is relatively low in moss taxa. No families are endemic to the páramo. Two genera are endemic, both described relatively recently. Kingiob1yum (Dicranaceae), originally described by Robinson (1967) and later in greater detail by Zander and Cleef ( 1982), is known from severallocalities in the Cordillera Oriental of Colombia and the Cordillera de Mérida of Venezuela. Gradsteinia (Amblystegiaceae ), an aquatic moss found on rocks in streams, is presently known only from the type locality in the Páramo de Sumapaz (Depto. Meta, Colombia) (Ochyra, 1990). At the species leve! approximately 9% are endemic; this figure is likely to be reduced after further taxonomic investigation and exploration, particularly in the central Andes. The concept of páramo adopted in our checklist for the mosses includes both subpáramo and páramo.
Elevation generally exceeds 3300 m, but in manyareas the páramo and páramo-like vegetation can extend down to 2800 m or less, and, at least for Colombia, páramo is frequently encountered around 3000 m. Defining the páramo and páramo plants (in this case, mosses) is not problematic when characterized by typical alpine-like open grassy vegetation, often dominated by members of the Poaceae such as Calamagrostis, Chusquea, and Festuca, cushion plants of various families and genera, and the often defining element Espeletia and related members of the tri be Espeletiinae for Venezuela south to northem Ecuador. However, challenges begin to arise when dealing with the forest elements present in the páramo zone. The first difficulty is related to the boundary between the montane forest and páramo; the subpáramo may be envisioned as the interfsce between the two zones, with open or shrubby montane forest grading into subpáramo or páramo. The second problem is the incursion of gallery forests into the páramo, and isolated forested escarpments within the páramo that harbor typical montan e mosses. It is only logical that the latter elements, gallery forest and forested escarpment, be included; but the former element,
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the interface between montane and páramo, is more subjective dueto the relative nature of zonations. The páramo zone (including subpáramo) exhibits a surprisingly wide range ofhabitats for mosses. The preponderance of species occur in exposed habitats on soil, humus, and rock, and many of these species readiiy ext¡:nd down slope into the open montane forest. Characteristic families of this group include the Bartramiaceae, Bryaceae, Dicranaceae, Ditrichaceae, Polytrichaceae, Pottiaceae, and Splachnaceae. A few families are generally restricted to the páramo, e.g., Andreaeaceae, Encalyptaceae, and Grimmiaceae (Racomitrium being the exception). Relatively few species are associated with streams and nearly always restricted to rocks, e.g., Andreaea nitida, A. acutifolia, Blindia magellanica, Dendrocryphaea latlfolia, Ditrichum bogotense, Fissidens rigidulus, Fon(inalis bogotensis, Holodontium strictum, Hygrodicranum bolivianum, Platyhypnidium aquaticum, and Schistidium rivulare. Typical forest elements, the majority found as epiphytes, include, among others, the Daltoniaceae, Macromitriaceae, Meteoriaceae, Neckeraceae, Orthotrichaceae, and Prionodontaceae. A few families---e.g., Brachytheciaceae, Hypnaceae, and Thuidiaceae-are found equally in exposed and forest si tes. Azonal habitats, such as marshy or boggy si tes, are characterized by such families as the Amblystegiaceae and Sphagnaceae. Severa! trends related to páramo mosses have been outlined by Griffin (1990a) for Venezuela, and they appearto be equally true for the whole ofthe páramo. The páramo zone is the second or possibly third most species-rich zone (for mosses) in the Neotropics. Given the extremely small surface area occupied by the páramo, it may even be the richest if species per area is considered in the assessment of diversity. The total number of species for the five tropical Andean countries is just over 2000 (Churchill et al., 1995). The northem Andean countries (Venezuela, Colombia, and Ecuador) are estimated to contain 1460 species (S. P. Churchill & D. Griffin, unpubl. data), of which approximately 34% of the species can be found in the páramo. The preponderance of páramo mosses (ca. 75%) are acrocarpic, tuft-forming mosses. Even among epiphytic mosses, a good number are acrocarpic or at least are erect in their growth form. Phytogeographical elements represented in the páramo include neotropical first, followed by Andean, cosmopolitan, Andean-African, neotropical-Asian, and finally, in equal parts, neotropical-Australasian, temperate Southem Hemisphere, and northern temperate/boreal. Studies related to páramo mosses include floristic inventaries (Bartram, 1953, 1964; Bowers, 1970;
Griffin, 1979; Magdefrau, 1983; Vareschi, 1970). Relatively few vegetational studies have included mosses-Balslev and de Vries (1991) did one, and of particular importance is the detailed study done by Cleef (1981 ). Although no floristic treatment of the páramo mosses presently exists, Churchill and Linares C. (1995) provide a synoptical flora ofColombia that includes páramo species, and D. Griffin is in the final stages of completing a full flora ofthe Venezuelan páramo. The following checklist has been compiled from the herbaria ofFLAS, NY, and US. Our own experiences in Costa Rica (DG), Venezuela (DG), Colombia (SPC & DG), and Ecuador (SPC) have also provided a certain amount of observational data for taxa both common and rare. The decision to include or exclude certain species, when confronted with marginal taxa or for which !abe! information was often inadequate, was based on our combined field knowledge. Literature has been employed only when no collections have been available in the above-mentioned herbaria. One hindrance in preparing this list related to labeled collections. At this time it seems incredible that professional systematists continue to provide inadequate !abe! information to accompany a collected specimen. Like sorne field-oriented taxonomists have done before us, we would urge workers to provide accurate !abe! information, including latitude and longitude, elevation ( or range ), associated vegetation type, and substrate. All taxa are listed in alphabetical order by family, genus, and species. Each country (six in al!) where a species is recorded as occurring in the páramo or páramolike vegetation (or at times marginal vegetation, as in subpáramo) is indicated. The elevational data, derived from a checklist in preparation for the tropical Andean countries, represent the entire elevation range ofthe species indicated only for Venezuela to Ecuador (in rare cases these have been modified due to additional data either from the highlands ofCentral America or Peru). For each species we have consulted voucher specimens from FLAS, NY, and US in an attempt to verify the species' occurrence in the páramo zone in each country. When we have not seen a specimen, the appropriate literature citation is given.
CHECKLIST OF MOSSES. ADELOTHECIACEAE Adelothecium Mitt. A. bogotense (Hampe) Mitt.; CO, VE;
1140~3300
m
55
PARAMOS
1999]
AMBLYSTEGIACEAE
BARTRAMIACEAE
Calliergon (Su11.) Kindb. C. luipichense R.S.Williams; EC; 4050 m C. sannentosum (Wah1enb.) Kindb.; VE, CO, EC; 35004050 m C. stramineum (Brid.) Kindb.; VE; 3100-4270 m C. trifarum (F.Weber & D.Mohr) Kindb.; VE; 3800 m
Anacolia Schimp. A. laevisphaera (Tay1or) F1owers; CO, VE, EC; 1850-4300 m
Cal/iergonel/a Loeske C. cuspidatum (Hedw.) Kindb.; CO, EC; 3000-4180 m Campyliadelphus (Lindb.) Broth. C. polyganus (Schimp.) Kanda; EC; 2000-3920 m Cratoneuron (Sull.) Spruce C.fi/icinum (Hedw.) Spruce; CO, VE, EC; 3000-3950 m Drepanocladus (Müll.Hal.) G.Roth D. aduncus (Hedw.) Wamst.; EC; 3800 m D. exannulatus (Schimp.) Wamst.; CO; 3450-3680 m D.jluitans (Hedw.) Wamst.; CO, VE, EC; 3000-4100 m D. leitensis (Mitt.) Broth.; EC; 2370-3800 m D. trichophyllus (Warnst.) Podp.; VE; 3700 m D. vernicosus (Mitt.) Wamst.; CO, VE; 3500-3800 m Gradsteinia Ochyra G. andicola Ochyra; CO; 3650 m Hygroamblystegium Loeske H. meridense E.B.Bartram; VE; 4000 m
Sanionia Loeske S. uncinata (Hedw.) Loeske; CO; 2800-4100 m
Sc01pidium (Schimp.) Limpr. S. cossonii (Schimp.) Hedenas; CO, EC; 3570-3950 m S. scorpioides (Hedw.) Limpr.; CO, VE, EC; 2900-4200 m S. turgescens (T.Jensen) Loekse; EC; 3800-4100 m
ANDREAEACEAE Acroschisma (Hook.f. & Wi1son) Lindl. A. wilsonii (Hook.f.) Lindl.; CO, EC; 3040-4100 m Andreaea Hedw. A. acutifolia Hook.f. & Wi1son; CO; 3700 m A. brevipes Spruce; CO, VE, EC; 3500-4765 m A. karsteniana Mü1l.Hal.; EC;? m [Note: Collected from the summit ofVo1cán Pichincha.] A. nítida Hook.f. & Wi1son; CO, VE, EC; 3240-4650 m A. rupestris Hedw.; CR, CO, VE, EC; 2900-4500 m A. seria/a G.Roth; EC; 5300 m A. striata Mitt.; EC; 5195 m A. urophylla H.Rob.; EC; 4200 m A. vulcanica Lorentz; EC; 3385-3690 m
AULACOMNIACEAE Aulacomnium Schwagr. A. palustre (Hedw.) Schwagr.; VE, EC; 3850-3950 m
Bartramia Hedw. Ref.: Fransén, 1995. B. angustifolia Mitt.; CO, VE, EC; 2800-4170 m B. brevifolia Brid.; CO, VE, EC; 2500-4030 m B. humilis Mitt.; CO, VE, EC; 1846-4000 m B. longifolia Hook.; CO, VE, EC; 2550-4070 m B. mathewsii Mitt.; CO, VE, EC; 3700-4360 m; inc1udes var. mathewsii and var. synoica Fransén B. polytrichoides Mü11.Hal.; CO, EC; 2620-4000 m B. potosica Mont.; CO, VE, EC; 2800-4765 m B. strumosa (Hampe) Mitt.; CR, CO, VE; 2800-4100 m Breutelia (Bruch & Schimp.) Schimp. Ref.: Griffin, 1988. B. brevifolia Herzog; CO; 3300-3745 m B. brittoniae Renau1d & Cardo!; CO, VE, EC; 2700-4030 m B. chrysea (Müll.Hal.) A.Jaeger; CO, VE, EC; 2200-4300 m B. inclina/a (Hampe & Lorentz) A.Jaeger; CO, VE, EC; 2100-3975 m B. integrifolia (Tay1or) A.Jaeger; CO, VE, EC; 3000-4150 m B. karsteniana (Müll.Hal.) A.Jaeger; CO, VE, EC; 25554200m B. maegdefraui H.A.Crum; CO; 3400 m B. polygastrica (Müll.Hal.) Broth.; CO, VE, EC; 2550-4300 m B. rhythidioides Herzog; CO, VE, EC; 2555-4000 m B. squarrosa A.Jaeger; CO, VE, EC; 2340-4000 m B. subarcuata (Müll.Hal.) Schimp.; CO, VE, EC; 1500-4300 m B. tomentosa (Brid.) A.Jaeger; CO, VE, EC; 1900-3930 m B. trianae (Hampe) A.Jaeger; CO, VE, EC; 2100-4340 m Conostomum Sw. C. tetragonum (Hedw.) Lindb. [syn.: C. pentastichum (Brid.) Lindb.); CR, CO, VE, EC; 3410-4200 m Flowersia D.G.Griffin & W.R.Buck F. setifolia (Hook. & Amott) D.G.Griffin & W.R.Buck; CO, PE; 4000-4300 m Leiomela (Mitt.) Broth. L. aristifolia (A.Jaeger) Wijk & Margad.; CO, EC; 32604250 m L. bartramioides (Hook.) Paris; CR, CO, VE, EC; 12003200 m L. deciduifolia Herzog; CO, EC; 2480-4250 m L. ecuadorensis H.Rob.; CO, EC; 3200-4200 m L. lopezii D.G.Griffin; CO, VE; 2440-3680 m L. peruviana R.S.Williams; EC; 3730-4155 m Philonotis Brid. P. andina (Mitt.) A.Jaeger; CO, VE, EC; 2770-4000 m P. cernua (Wilson) D.G.Griffin & W.R.Buck; VE; 3300 m P. crassinervia Broth. & Paris; VE; 1700-3600 m P. erecta (Mitt.) D.G.Griffin & W.R.Buck; VE; 3200 m P.fontanella (Hampe) A.Jaeger; CO, EC; 2700-3750 m P. has/ata (Duby) Wijk & Margad.; VE; 2900-3500 m
·' 1
MEMOIRS OF THE NEW YÜRK BOTANICAL GARDEN
56
P. /ongiseta (Michx.) E.Britton; CR, 3300 m P. revoluta Bosch & Sande Lac.; CO; 2100--4100 m P. rufzjlora (Homsch.) Reichardt; CO, VE, EC; 1000--4000 m P. scabrifolia (Hook.f. & Wilson) Braithw.; CO, VE, EC; 3750-4170m
BRACHYTHECIACEAE Aero/indigia M.Menzel A. capil/acea (Hornsch.) M.Menzel; CO, EC; 700-3700 m Brachythecium Schimp. Ref.: McFarland, 1988. B. austro-glareosum (Müli.Hal.) Kindb.; CO, EC; 33004100 m B. chocayae Herzog; CO; 3550 m B. cirriphylloides McFarland; CO, VE; 2900-3620 m B. conostomum (Taylor) A.Jaeger; CO, VE, EC; 3100--4500 m B. accidenta/e (Hampe) A.Jaeger; CO, VE, EC; 2500--4500 m B. plumosum (Hedw.) Schimp.; CO, VE, EC:; 2000-3600 m B. praelongum Schimp. ex Müll.Hal.; CO, VE, EC; 36004350 m B. rutabulum (Hedw.) Schimp.; CO, VE, EC; 2400--4050 m B. stereopoma (Mitt.) A.Jaeger; CO, VE, EC; 900-3650 m Eurhynchium Schimp. E. prae/ongum (Hedw.) Schimp.; CR, VE, EC; 2100-3780 m Palamocladium Müii.Hal. P. leskeoides (Hook.) E.Britton; CO, VE, EC; 2000-3300 m Platyhypnidium M.Fleisch. P. aquaticum (A.Jaeger) M.F1eisch.; CO, VE, EC; 15004200 m Rhynchostegium Bruch & Schimp. R. inerme (Mitt.) A.Jaeger; EC; 3810 m R. scariosus (Taylor) A.Jaeger; CO, VE, EC; 800-3680 m R. serrulatum (Hedw.) A.Jaeger; CO, VE, EC; 1000--4000 m Rozea Be'sch. R. chrysea B~sch.; VE; 3500-3520 m. R. subjulacea B~sch.; CO VE; 3300--3670 m
BRUCHIACEAE Eobruchia W.R.Buck E. ecuatoriana Steere; EC; 4200 m
BRYACEAE Ref.: Ochi, 1980, 1981. Acidodontium Schwagr. A. heteroneuron (Mitt.) Broth.; VE, EC; 1600-3750 m A. /ongifolium (Paris) Broth.; EC; 3690 m. A. mega/ocarpum (Hook.) Renau1d & Cardo!; CO, EC; 2310--4050 m A. seminerve Hook.f. & Wilson; CO, EC; 1540-3825 m A. subrotundum (Tay1or) Hook.f. & Wilson; EC; 3400 m
[VOL. 84
, Anomobtyum Schimp. A. clavicaule (Müll.Hal.) Broth.; CO, VE; 3085-3965 m A. conicuin (Homsch.) Broth.; EC; 1050--4000 m A. ju/aceum (P.Gaertn., B.Mey. & Scherb.) Schimp. (syn.: A.filiforme (Dicks.) Solms.]; CR, CO, VE, EC; 23204050 m A. p/icatum Cardo!; CR, CO, VE; 3300-3700 m A. prostratum (Müll.Hal.) Besch.; CO, VE, EC; 1230-3890 m A. robu'stum Broth.; CO; 3000-3525 m Brachymenium Schwagr. B. globosum A.Jaeger; EC; 1200-3450 m B. speciosum (Hook.f. & Wi1son) Steere; CO, EC; 12303385 m B. systylium (Mü11.Hal.) A.Jaeger; VE, EC; 3020-3600 m Btyum Hedw. B. a/govicum Sendtn. ex Mü1l.Hal.; VE, EC?; 3800--4400 m B. alpinum Hüds. ex With.; CO, VE; 2900-3700 m B. andicola Hook.; CR, CO, VE, EC; 1100--4325 m B. argentelfm Hedw.; CR, CO, VE, EC; 20--4250 m B. biforme R.S.Williams; VE; 4070 m B. canariense Brid.; CO; 3620 m B. capil/are Hedw.; CR, CO, VE, EC; 2285--4260 m B. chryseum Mitt.; CR; 2200-3600 m B. densifo/ium Brid.; EC; 940-3385 m B. dichotomum Hedw.; CO, EC; 1050--4100 m B. ellipsifo/ium Müll.Hal.; VE, CO; 3630--4400 m B. goudotii Hampe; VE; 3350-3600 m B. incacorralis Herzog; VE; 4070 m B. innovans H.A.Crum; CO; 4150 m B. /aevigatum Hook.f. & Wilson; CO, VÉ; 3000--4335 m B. micro-nitidum Ochi; EC; 375'0-3860 m B. pal/escens Schleich. ex Schwagr.; CO, EC; 3080--4340 m B. pseudotriquetrum (Hedw.) P.Gaertn., G.Mey. & Scherb.; CO, VE, EC; 1400--4100 m LeptobtJ'llm (Bruch & Schimp.) Wi1son L. pyriforme (Hedw.) Wilson; CO, VE, EC, PE; 2750--4100 m L. wilsonii (Mitt.) Broth.; EC; 3700 m Mielichhoferia Nees, Hornsch. & J.W.Sturm M. antisanensis (E.B.Bartram) H.Rob.; CO, EC; 3440 m ,M, argentifolia Mitt.; VE, EC; 3900 m M. dip/odonta Mitt.; EC; 3110-3505 m M. laxiretis Thér.; EC; 4000--4400 m M. longiseta Mü1l.Hal.; EC; 2400-5110 m M megalocmpa (Amott) A.Jaeger; CO, VE, EC; 2215--4000 m Orthodontium Schwagr. O. graci/e Schwagr. ex Bruch, Schimp. & W.Gümbel; VE; 3000-3400 m O. pellucens (Hook.) Bruch, Schimp. & W.Gümbel; CR, CO, VE, EC; 1420-3400 m Pohlia Hedw. P. chilensis (Mont.) A.J.Shaw; EC; 3500-3900 m P. elongata Hedw.; CO, VE, EC; 1450-3450 m P. /ongicollis (Hedw.) Lindb.; VE; 3100-3250 m
/
1999]
P. P. P. P.
PARAMOS
papillosa (A.Jaeger) Broth.; CO, VE, EC; 2200-4045 m richardsii A.J.Shaw; CR; 3100 m schisticola (Müll.Hal.) Broth.; PE; 2700-3100 m wahlenbergii (F.Weber & D.Mohr) A.L.Andrews; CO, VE, EC; 2900-4045 m
Rhodob1yum Hampe R. beyrichianum (Homsch.) Müll.Hal.; VE; 1300-3600 m R. grandifolium (Tay1or) Schimp.; CO, VE, EC; 1300-4000 m R. perspinidens (Broth.) Pócs; CO; 3220-3780 m R. procerum (Besch.) París; CO, VE, EC; 2900-3700 m R. roseodens (Mü1l.Hal.) Paris; CO, VE; 3350-4000 m Schizymenium Harv. S. andinum (Sull.) A.J.Shaw; VE; 2500-4000 m S. bogotense (Hampe) A.J.Shaw; CO, EC; 2700-3800 m S. campylocarpum (Hook. & Amott) A.J.Shaw; CR, CO, VE, EC; 3300-4100 m S. fusiferum (Mítt.) A.J.Shaw; CO, EC; 2925-3800 m S. gracilisetum (Hampe) A.J.Shaw; CO, VE; 2200-3600 m S. lindigii (Hampe) A.J.Shaw; CO, EC; 2600-3700 m S. linearicaulis (Mü1l.Hal.) A.J.Shaw; EC; 3910 m S. lonchocarpum (Hook. & Amott) A.J.Shaw; VE; 3390 m S. nanum Tay1or; EC; 4115-4570 m . S. pseudopohlia A.J.Shaw; CO, VE, EC; 2900-3800 m S. pusillum (Hook.f. & Wilson) A.J.Shaw; VE; 3100-3600 m
CALLICOSTACEAE Callicostellopsis Broth. C. meridensis (Mü11.Hal.) Broth.; VE; 3400-3620 m Cyclodicyton Mítt. C. roridum (Hampe) Kuntze; CO, EC; 1950-3950 m Lepidopilum (Bríd.) Brid. L. longifolium Hampe; CO, VE, EC; 1790-3700 m Trachyxiphium W.R.Buck T. steerei (D.G.Griffin) S.P.Churchill; VE, CO; 3500-3755 m
CATAGONIACEAE Catagonium Mü11.Hal. C. brevicaudatum Müii.Hal. ex Broth.; CO, VE, EC; 27004250 m
CRYPHAEACEAE Crypháea D.Mohr C.fasciculosa Mitt.; VE; 2900-3600 m C. jamesonii Taylor; VE, EC; 1850-3900 m C. patens Homsch.; CO, EC; 1520-3700 m C. pilifera Mítt.; CO, VE, EC; 1800-3500 m Dendrocryphaea París & Schimp. ex Broth. D. latifolia D.G.Gríffin, Gradst. & J.Aguirre; CO; 30803400m
57
DALTONIACEAE Daltonia Hook. & Tay1or Re f.: Bartram, 1931. D. bilimbata Hampe; CO, VE, EC; 2000-4100 m D. brevinervis E.B.Bartram; CO; 3300 m D.fenestrellata D.G.Griffin; CO; 3350 m D. gracilis Mítt.; CR, CO, VE, EC; 2100-4200 m D. jamesonii Taylor; CO, EC; 3000-4260 m D. latolimbata Broth.; VE; 2900-3500 m D. lindigiana Hampe; CR, CO, EC; 2620-4000 m D. longifolia Tay1or; CO?, EC; 1540-4080 m D. peruviana Mitt.; VE, EC; 3400-4100 m D. pulvinata Mitt.; CR, CO, EC; 1800-4200 m D. stenophylla Mitt.; CO, EC; 3000-3850 m D. tenuifolia Mitt.; CO?, EC; 2050-3500 m D. t1'achyodonta Mítt.; VE, EC; 3400-4000 m Leskeodon Broth. L. andicola (Mitt.) Broth.; CO, EC; 310-3730 m
DICRANACEAE Ref.: Frahm, 1991.
Aongstroemia Bruch & Schimp . A.julacea (Hook.) Mitt.; CR, CO, VE, EC; 3000-4600 m A. orientalis Mítt.; VE; 2900-3800 m Campylopus Bríd. C. albidovirens Herzog; CO, VE, EC; 2500-4000 m C. amboroensis Thér.; VE?, CO; 3575-4300 m C. anderssonii (Müll.Hal.) A.Jaeger; CO; 3260-4100 m C. arctocarpus (Homsch.) Mítt.; CO, VE?, EC; 2200-3700 m C. areodictyon (Mü11.Hal.) Mitt.; CO, VE, EC; 2900-3950 m C. argyrocaulon (Mü11.Hal.) Broth.; CO, VE, EC; 30004000 m C. asperifolius Mítt.; CO, VE, EC; 2000-3350 m C. bryotropii J.-P.Frahm; VE, CO; 3500-3730 m C. capitulatus E.B.Bartram; VE, EC; 2000-3950 m C. cavifolius Mitt.; CO, VE?, EC; 3240-4300 m C. cleefii J.-P.Frahm; CO; 3300-4400 m C. densicoma (Müll.Hal.) París; VE, CO; 2100-4250 m; includes var. densicoma and var.yungarum (Herzog) J.P.Frahm C. dicnemioides (Mü11.Hal.) París [syn.: C. cucullatifolius Herzog, C. cuspidatus (Homsch.) Mitt. var. dicnemioides (Mü11.Hal.) J.-P.Frahm]; CO, VE, EC; 2450-3950 m C. edithae Broth.; CO, VE, EC; 3600-4100 m C. flexuosus (Hedw.) Brid.; CR, CO, VE, EC; 2850-3950 m; íncludes var..flexuosus and var. incacorralis (Herzog) J.-P.Frahm C. fragilis (Bríd.) Bruch & Schimp. var. fragilis; CR, CO, VE?, EC; 1550-4000 m C. heterostachys (Hampe) A.Jaeger; CO, VE, EC; 16404300m C. huallagensis Broth.; CO, EC; 1800-4100 m C. incertus Thér.; CO, VE, EC; 2800-3850 m C.jamesonii (Hook.) A.Jaeger; CR, CO, VE, EC; 2180-4000 m
58
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
C.jugorum Herzog; CO; 3100-4235 m C. lamellinervis (Müll.Hal.) Mitt. var. /amellinervis; CO;
3300-3520 m C. longicel/ularis J.-P.Frahm; VE?, CO; 2970-3830 m C. nivalis (Brid.) Brid.; CR, CO, VE, EC; 2500-4800 m; includes var. multicapsularis (Mü1l.Hal.) J.-P.Frahm and var. nivalis C. oblongus Thér.; VE, CO; 2900-3200 m C. pauper (Hampe) Mitt. var. pauper; CO, EC; 2200-3400 m C. perexilis (Müll.Hal.) Paris; CO; 3750 m C. pilifer Brid. var. pilifer; CO?, VE, EC; 1500-4000 m C. pittieri R.S.Williams; CO, VE, EC; 2500-4200 m C. rejlexisetus (Müll.Hal.) Broth.; CO, VE, EC; 2600-3690 m C. richardíi Brid.; CR, CO, VE, EC; 600-3910 m C. sharpii J.-P.Frahm, D.G.Horton & Vitt; CO, EC; 31Q03570 m C. subcuspidatus (Hampe) A.Jaeger; CO, EC; 3650-3850 m C. subjugorum Broth.; CO, VE, EC; 3050-4250 m C. thysanomitrioides Thér.; VE; 3600-3750 m C. trichophylloides Thér.; CO, EC; 3850-4700 m C. trivialis Mü11.Hal. ex E.Britton; CO, EC; 1600-4300 m Chorisodontíum (Mitt.) Broth.
Ref.: Frahm, 1989. C. mittenií (Müli.Hal.) Broth.; CO, VE, EC; 2100-3750 m C. wallisii (Müli.Hal.) Broth.; CO, VE, EC; 2340-4050 m; includes var. speciosum (Hook.f. & Wilson) Broth. and var. wallisii Dicranella (Müll.Hal.) Schimp. [Note: Includes Anisothecíum Mitt.] D. campylophyllum (Tay1or) A.Jaeger [syn.: Cheilothela vaginata H.Rob.; D. cardotii (R.Br. ter.) Dixon]; CO,
VE, EC; 2940-4400 m D. hookeri (Müli.Hal.) Cardot; CO, VE, EC; 2240-4100 m D. vaginatum (Hook.) Cardot; CO, EC; 2700-3950 m Dicranodontíum Bruch & Schim¡i. D. pulchroalare Broth. (syn.: D. meridiana/e E.B.Bartram);
CO; 2660-3500 m Dicranoweisia Lindb. ex Milde D.fastigiata (Mitt.) Paris; EC; 3000-4410 m Dicranum Hedw. D.frigídum Müli.Hal.; CR, CO, VE 1 EC; 1600-3800 m D. peruvianum H.Rob.; CO, EC, PE?; 3200-3800 m Holodontíum (Mitt.) Broth. H. strictum (Hook.f. & Wilson) Ochyra [syn.: H. inerme
[VOL. 84
Kingiobryum H.Rob. K. paramicola H.Rob.; VE, CO; 3490-4200 m Metzleria Schimp. ex Milde (syn.: Atracty/ocarpus Mitt.) M. longiseta (Hook.) Broth.; CR, CO, VE, EC; 2000-4300 m Oreoweisia (Bruch, Schimp. & W.Gümbel) De Not.
Ref.: Griffin, 1986. O. O. O. O.
brasiliensís Hampe;VE, CO; 2600-3200 m erosa (Müll.Hal.) Kindb.; CO, VE, EC; 2600-4500 m laxiretís Broth. ex Herzog; VE; 3390 m tunariensís Herzog; VE, PE; 3100-3900 m
Pilopogon Brid. P.- guade!oupensis (Brid.) J.-P.Frahm; CR, CO, VE, EC;
1440-4400 m P. laevis (Taylor) Thér.; CO, VE, EC; 3200-4410 m P. macrocarpus Broth.; CO; 2700-3500 m Rhabdoweisia Bruch & Schimp. R.fugax(Hedw.) Bruch & Schimp.; CR, CO,EC; 2200-3600 m Sphaerothecíum Hampe S. phascoides Thér.; CO; 2770-3400 m Symblepharís Mont. S. fragilis Mitt.; EC; 2770-4000 m S. líndigií Hampe; CO, EC; 2500-3750 m S. vaginata (Hook.) Wijk & Margad.; EC; 1850-3690 m
DITRICHACEAE Ceratodon Brid. C. stenocarpus Bruch & Schimp.; CR, CO, VE, PE; 2400-
3900 m Chrysoblastella R.S.Williams C. chilensís (Mont.) Reimers; VE, CO, EC; 2900-4200 m Distichum Bruch & Schimp., nom. cons. D. capillaceum (Hedw.) Bruch & Schimp.; CO, VE, EC;
3500-4280 m Ditrichum Hampe, nom. cons. D. erina/e (Tay1or) Kuntze; CO, VE, EC; 2700-4300 m D. gracile (Mitt.) Kuntze; CO, EC; 3000-4350 m D. rufescens (Hampe) Hampe; CO, VE, EC; 2000-3700 m D. submersum Cardot & Herzog; CO; 3600-3800 m D. bogotense (Hampe) Broth. (syn.: D. venezuelanum
D.G.Griffin); VE, CO; 3100-3240 m
(Mitt.) Broth.]; EC; 4200 m Holomitrium Brid. H. arboreum Mitt.; CO, EC; 500-3500 m H.jlexuosum Mitt.; CO, EC; 1800-3500 m H. pulchellum Mitt.; CR, CO, EC; 2460-4000 m Hygrodicranum Cardo! H. bo!ivíanum Herzog; EC; 3900-4100 m
Pleurídium Rabenh. P. andínum Herzog; VE; 3500 m P. lindígianíum (Hampe) S.P.Churchill; CO; 2500-3200 m P. venezuelanum D.G.Griffin; VE; 3800 m Trístichíum Müll.Hal. T. lorentzií Herzog; CO; 4130 m T. mirabile (Müll.Hal.) Herzog; CR, CO, VE; 3300-3600 m
1999]
PARAMOS
ENCALYPTACEAE
59
GRIMMIACEAE
EUSTICHIACEAE
Grimmia Redw. G. donniana Sm.; EC; 4650 m G.fuscolutea Rook.; VE, EC; 3960--4650 m G. /ongirostris Rook.; CR, CO, VE, EC; 2800-4400 m [Note: G. affinis Romsch. and G. ovalis (Hedw.) Lindb. ofreports.] G. navicularis Rerzog; CO, VE, EC; 3850-5300 m G. ova lis (Hedw.) Lindb.; CR, VE, EC; 3100--4060 m G. pansa R.S.Williams; CO, VE, EC; 3200--4400 m G. trichophylla Grev.; CR, PA, CO, VE, EC; 2900-3900 m
Diplostichum Mont. (syn.: Eustichia Broth.) D. longirostre (Brid.) Brid.; CR, CO, VE, EC; 2200--4030 m
Ptychomitrium Fürnr. P. chimborazense (Mitt.) A.Jaeger; VE, EC?; 2615-3200 m
FABRONIACEAE
Racomitrium Brid. R. brachypus Mü1l.Ral.; EC; 3700 m R. crispipilum (Tay1or) A.Jaeger; CR, CO, VE, EC, PE; 2600--4500 m R. cucullatifolium Rampe; VE, CO, EC; 2400-3885 m R. dichelymoides Herzog; CO; 1500-3800 m R. lanuginosum (Hedw.) Brid.; CO, EC, PE; 3000--4200 m R. steerei D.G.Griffin; VE; 3370-3750 m
Encá!ypta Hedw. E. asperifolia Mitt.; CO, VE, EC; 3600--431 O m E. ciliata Hedw.; CO, EC; 3080--4200 m
ENTODONTACEAE Entodon Müll.Hal. E.jamesonii (Taylor) Mitt.; CO, VE, EC; 1000-3500 m
Fabronia Raddi F. ciliaris (Brid.) Brid. var. wrightii (Su!!.) W.R.Buck; EC; 1800-3500 m F. jamesonii Taylor; VE; 3100-3200 m
FISSIDENTACEAE Fissidens Í-!edw. F. crispus Mont. (syn.: F. repandus Wilson ex Mitt.); CO, F. F. F. F.
EC; 200-3500 m curvcllus Homsch. (syn.: F. milobakeri L.F.Koch); VE; 2900-3500 m rigidulus Hook.f. & Wilson; CO, VE, EC; 2200--4100 m serrulatus Müll.Ral. var. leptochaete (Dusén) Brugg.Nann. & Pursell; EC; 3700 m wallisii Müll.Ral.; CO; 2200-3700 m
Schistidium Brid. S. angustifolium (Mitt.) Herzog; EC; >4000 m S. apocarpum (Redw.) Bruch & Schimp.; CO, VE, EC; 2900-3890 m S. rivulare (Brid.) Podp.; CO, VE, EC; 3530--4000 m; ineludes subsp. /atifolium (Brid.) Podp. in EC and subsp. rivulare
HEDWIGIACEAE FONTINALIACEAE Fontinalis Redw. F. bogotensis Hampe; CO; 2000-3700 m
FUNARIACEAE Entosthodon Schwiigr. Ref.: Fife, 1987. E. bonplandii (Hook.) Mitt.; CR, CO, VE, EC; 1545-3690 m E.jamesonii (Taylor) Mitt.; CO, VE, EC; 2200--4200 m E. /aevis (Mitt.) Fife; EC; 2000-3900 m E. /axus (Wilson & Hook.f) Mitt.; VE, EC; 3400--4100 m E. lindigii (Rampe) Mitt.; CO, VE, EC; 2500-3690 m E. mathewsii Hook.f.; EC; 3385 m E. obtusifolius Rook.f.; VE; 1850-3500 m E. radians (Redw.) Müll.Ral. [syn.: E. acaulis (Hampe) Fife]; VE, EC; 1900-3310 m Funaria Redw. F. ca!vescens Schwiigr.; CR, CO, EC; 1000-3 750 m F. hygrometrica Redw.; CO, VE; 2900--4400 m F. macrospora R.S.Williams; VE; 4000 m F. microstoma Bruch ex Schimp.; VE; 2900-3300 m
Braunia Bruch & Schimp. B. cirrhifolia (Wilson) A.Jaeger; EC; 3000-3300 m B. diaphana (Müll.Ral.) A.Jaeger [syn.: B. plicata (Mitt.) A.Jaeger]; EC; 3200-3300 m B. /axifolia Rerzog; VE; 3100-3200 m B. nephelogenes DeLuna & W.R.Buck; EC; 3600-3755 m B. secunda (Rook.) Bruch & Schimp.; VE, EC; 3100--4150 m Hedwigia P.Beauv. H. ciliata (Redw.) P.Beauv.; CO, VE, EC; 1800-3690 m Hedwigidium Bruch & Schimp. H. integrifolium (P.Beauv.) Díxon; CR, CO, VE, EC; 24204150 m
HYLOCOMIACEAE Loeskeobryum M.Fieísch. ex Broth. L. giganteum (E.B.Bartram) J.R.Rohrer; CO; 3570 m Pleurozium Mitt. P. schreberi (Brid.) Mitt.; CO, VE?, EC; 2650--4200 m
60
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
Rhytidium (Sull.) Kindb. R. rugosum (Hedw.) Kindb.; VE; 3100 m
HYPNACEAE Cariobaeohypnum Ando & Higuchi C. po/ypterum (Mitt.) Ando & Higuchi [syn.: Hypnum polypterum (Mitt.) Broth.]; CO, VE, EC;2800-3500 m Ctenidium (Schimp.) Mitt. C. malacodes Mitt.; CO, VE, EC; 1700-4000 m Herzogiel/a Broth. H cy/indricarpa (Cardot) Z.lwats.; CO, VE, EC; 2600-4100 m Hypnum Hedw., noin. cons. H. amabi]e (Mitt.) Hampe; CR, CO, VE, EC; 1850-3750 m H. cupressiforme Hedw.; CO, VE, EC; 2900-4100 m; ineludes var.lacunosum (Brid.) Bertsch. and var. cupressiformein CO
MACROMITRÍACEAE Macrocoma (Mü11.Hal.) Grout M. tenue (Hook. & Grev.) Vitt subsp. sullivantii (Mü1l.Hal.) Vitt; VE, EC; 1400-3300 m Macromitrium Brid. M. aureum Mü11.Hal.; CO, EC; 2460-3750 m M. cirrosum (Hedw.) Brid.; CR, CO, EC; 520-4200 m M. drewii H.Rob.; EC; 3750-3850 m M.fiiscoaureum E.B.Bartram; CO, VE, EC; 1850-3385 m M. guatemalense Mü11.Hal. (syn.: M. serrulatum Mitt.); CO, VE, EC; 1940-3750 m M. longifolium (Hook.) Brid.; CO, VE, EC; 1800-4220 m M ob/ongum (Tay1or) Mitt.; EC; 3100 m · Ú. perrejlexum Steere; EC; 2750-3700 m M. punctatum (Hook. & Grev.) Brid.; EC; 300-3300 m ·M. subcirrosum Müll.Hal.; CO; 1840-3550 m M. subcrenulatum Broth.; EC; 3200-3600 m M. trachypodium Mitt.; CO, VE, EC; 1800-3630 m M. ulophyllum Mitt.; CO, EC; 2000-3600 m
Mittenothamnium Henn., nom. cons. M. /ehmannii (Besch.) Cardot; CR; 1300-3500 m M. reptans (Hedw.) Cardot; CO, VE, EC; 975-3400 m Pseudotaxiphyllum Z.Iwats. P. distichaceum (Mitt.) Z.Iwats.; CO, VE; 3700 m
[VOL. 84
MEESIACEAE Meesia Hedw. M. longiseta Hedw.; CO, EC; 2300-3800 m M. triquetra (Hedw.) Angstr.; VE; 3500 m
Pylaisiel/a Kindb. ex Grout P.fa/cata (Schimp.) Ando; CR, CO, VE, EC; 2460-4000 m
METEORIACEAE Lindigia Hampe
HYPOPTERYGIACEAE Hypopterygium Brid. H. tamarisci (Sw.) Brid. ex Müll.Hal.; VE, EC; 900-3300 m
L. debi/is (Mitt.) A.Jaeger; CO, VE, EC; 2580-3400 m
Meteorium (Brid.) Dozy & Mo1k. M. sinuatum (Müll.Hal.) Mitt. (syn.: M. il/ecebrum Sul1.); CO, VE, EC; 1000-3775 m
LEPYRODONTACEAE Lepyrodon Hampe L. tomentosus (Hook.) Mitt.; CR, PA, CO, VE, EC; 29004300 m
LESKEACEAE Leskea Hedw. L. angustata Tay1or; Vp, EC; 2155-3500 m L. teretiuscu/a Mitt.; EC; 2460-3385 m Leskeadelphus Herzog L. bolivianus (E.Britton) W.R.Buck; CO, EC; 2300-3500 m
LEUCODONTACEAE Leucodon Schwagr. L. curvirostris Hampe; CR, VE, CO; 2600-3600 m
Papillaria (Müll.Hal.) Mü11.Hal. P. deppei (Homsch.) A.Jaeger; CO, VE; 1030-3500 m P. impon de rosa (Taylor) Broth.; CO, VE, EC; 130-3100 m P. penici/lata (Dozy & Molk.) Broth.; VE; 1300-3000 m Pilotrichella (Müll.Hal.) Besch. P. jlexilis (Hedw.) Ángstr.; CO, VE, EC; 1725-3550 m P. quitensjs (Mitt.) A.Jaeger; EC; 2850-3500 m Squamidium (Müll.Hal.) Broth. S. /eucotrichum (Taylor) Broth.; CR, CO, VE?, EC; 6003800m S. nigricans (Hook.) Broth.; CO, EC; 750-3500 m Zelometeorium Manuel Z. patulum (Hedw.) Manuel; VE; 20-3200 m
MNIACEAE LEUCOMIACEAE Rhychostegiopsis Mü1l.Hal. R. tunguraguana (Mitt.) Broth.; CO, EC; 2600-3450 m
Plagiomnium T.J.Kop. P. rhynchophorum (Hook.) T.J.Kop.; CR, CO, VE, EC; 5003850m
1999]
PARAMOS
NECKERACEAE Neckera Hedw. andina Mitt.; EC; 3500 m chilensis Schimp. ex Mont.; CO, VE, EC; 1680-4100 m ehrenbergiiMüll.Hal.; CO, EC; 3100-4100 m obtusifolius Taylor; CO, VE, EC; 2000-4000 m scabridens Müll.Hal.; CO, EC; 1700-3750 m
N. N. N. N. N.
ORTHOTRICHACEAE Amphidium Schimp., nom. cons. A. tortuosum (Hornsch.) Cufod. [syn.: A. cyathicarpum (Mont.) Broth.]; CO, VE, EC; 3000-4100 m Orthotrichum Hedw. Ref.: Lewinsky, 1984, 1987. O. aequatoreum Mitt.; VE, EC; 2200-3800 m O. elongatum Taylor; CO, VE, EC; 2560-4050 m O. latimarginatum Lewinsky; EC; 3780 m O. laxifolium Wilson; VE, EC; 2900-3690 m O. mandonii Schimp. ex Hampe; CO, VE, EC; 3000-4120 m O. pallens Brid.; VE; 3100-3300 m O. pariatum Mitt.; CR, CO, VE, EC; 2590-4000 m O. penicillatum Mitt.; CO, VE; 2700-4000 m O. pungens Mitt.; EC; 2600-4000 m O. pycnophyllum Schimp.; CO, VE, EC; 2200-4100 m; ineludes var. pycnophyllum and var. verrucosum Lewinsky O. rupestre Schleich. ex Schwagr.; VE, EC; 3200-4210 m O. steerei Lewinsky; CO, VE, EC; 3200-3750 m O. subulatum Mitt.; EC; 3100-3200 m O. trachymitrium Mitt.; EC; 3500 m Zygodon Hook. & Taylor Ref.: Griffin, 1990b; Malta, 1926. Z. altarensis Broth.; EC; 4000-4200 m Z. campylophyllus Müll.Hal.; CO; 3550 m Z. ehrenbergii Müll.Hal.; CO, EC; 2000-3500 m Z.fasciculatus Mitt.; EC; 3000-4100 m Z.fi'agilis H.Rob.; CO, VE, EC; 3220-4200 m ,z. goudotii Hampe; VE; 2700-3800 m Z. liebmannii Schimp.; CR, EC; 3200-3750 m Z. longicel/ularis D.G.Griffin; VE; 3600-3750 m Z. nivalis Hampe; CO, EC; 3300-4050 m Z. oeneus Herzog; VE; 3200 m Z. peruvianus Sull.; CO, VE, EC; 2460-4230 m Z. petrophilus R.S.Williams; VE; 2900-3300 m Z. pichinchensis (Taylor) Mitt.; CO, VE, EC; 3300-451 Om Z. quitensis Mitt. var. integrifolius Malta; EC; 4000-4200 m Z. reinwardtii (Hornsch.) Braun; CR, CO, VE, EC; 19403750m Z. rufescens (Hampe) Broth.; VE; 2900-3600 m Z. sordidus Müll.Hal.; VE; 2900-3300 m Z. squarrosus (Taylor) Müll.Hal.; CO, EC; 3000-4200 m Z. stenocarpus Taylor; CO, EC; 3200-4100 m; includes var. !inearifolius (Mitt.) Malta and var. stenocarpus Z. subsquarrosus Broth.; EC; 3810-4000 m Z. venezuelensis D.G.Griffin; VE; 3700 m
61
PHYLLOGONIACEAE Phyllogonium Brid. P. fu/gens (Hedw.) Brid.; CO, EC; 500-4200 m P. viscosum (P.Beauv.) Mitt.; CO, EC; 1000-3500 m
PLAGIOTHECIACEAE Plagiothecium Bruch & Schimp. P. conostegium Herzog; CO, VE, EC; 3000-4040 m P. drepanophyllum Renau1d & Cardot; CO, VE; 2500--3740 m P. lucidum (Hook.f. & Wilson) Paris; CO, VE, EC; 20004120m P. novo-granatense (Hampe) Mitt.; CO, EC; 2600-3700 m
POLYTRICHACEAE Notoligotrichum G.L.Sm. N. trichodon (Hook.f. & Wilson) G.L.Sm.; CO, EC; 40004200 m 0/igotrichum Lam. & A.DC. O. erosum (Hampe) Lindb.; CO?, EC; 2600-3400 m Pogonatum P.Beauv. Ref.: Hyviinen, 1989. P. campy/ocarpum (Müll.Hal.) Mitt.; CR, CO, VE, EC; 1400-4000 m P. neglectum (Hampe) A.Jaeger; CO, VE, EC?; 1900--3600 m P. perichaetiale (Mont.) A.Jaeger var. oligodus (Müll.Hal.) Hyviinen; CO, VE, EC; 1700-4180 m P. procerum (Lindb.) Schimp.; EC; 3600 m Polytrichadelphus (Müll.Hal.) Mitt. P. aristatus (Hampe) Mitt.; CO, VE, EC; 1140-4200 m P. ciliatus (Hook.f. & Wilson) Mitt.; CO, VE; 2500--3500 m P. costaricensis E.B.Bartram; CR; ? m P. giganteus (Hook.) Mitt.; CO; 3500 m P. /ongisetus (Hook.) Mitt.; CO, VE, EC; 1440-4080 m P. purpureus Mitt.; CO, EC; 3000-3900 m Polytrichastrum G.L.Sm. P. alpiniforme (Cardot) G.L.Sm.; CR, EC; 3500-4120 m P. tenel/um (Müll.Hal.) G.L.Sm.; CO, VE; 2800-4100 m Polytrichum Hedw. P. ericoides Hampe; CO, EC?; 1500-3450 m P.juniperinum Hedw.; CR, CO, VE, EC; 800-4680 m
POTTIACEAE Ref.: Zander, 1993.
Alonia Kindb. A. rigida (Hedw.) Limpr.; EC; 3385 m Aloinella Cardot A. cucullatifolia (Müll.Hal.) Broth.; EC; 3690 m A. cucul/ifera (Mü1l.Hal.) Steere; EC; 3385-4310 m A. venezue/ana D.G.Griffin; VE; 3800-4000 m
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MEMOIRS OF TRE NEW YORK BOTANICAL GARDEN
L. longicau/e Mitt.; CR, CO, VE, EC; 2350-4600 m; ineludes var. /ongicaule and microruncinatum (Dusén·)
Anoectangium Schwagr. A. aestivum Redw. [syn.: A. euchloron (Schwagr.) Mitt.];
R.R.Zander
CO, VE, EC; 1120-4030 m Barbula Redw., nom. cons. B. appressifolia (Mitt.) A.Jaeger; CO; 4100 m B. costa/a (Mitt.) A.Jaeger; EC; 2200-4100 m B. glaucescens Rampe; CO, EC; 3200-4000 m B. vulcanica Lorentz; EC; 2770-3690 m Bellibarbula P.C.Chen B. recurva (Gdff.) R.R.Zander [syn.: B¡yoerythrophyllum recurvum (Griff.) K.Saito]; VE; 3200 m Bryoerythrophyllum ~.C.Chen B. bolivianum (Mü1l.Ral.) R.R.Zander; EC; 3690 m B. campylocarpum .(Müli.Ral.) R.A.Crum; CO, VE, EC;
2100-3500 m B. inaequalifolium (Tay1or) R.R.Zander; CO, VE, EC; 3000-
4300 m B.jamesonii (Tay1or) R.A.Crum; CO, VE, EC; 1400-4300 m Didymodon Redw. D. australasiae (Roo k. & Grev.) R.R.Zander; CO, VE, EC;
1650-4300 m D. inundatus (Mitt.) Broth.; EC; 3570 m D. laevigatus (Mitt.) R.R.Zander; CO, VE, EC; 2400-4350 m D. nigrescens (Mitt.) Saito (syn.: Barbula brunneola Müli.Ral.); CR; 3491 m D. rigidulus Redw.; VE, EC; 1700-4300 m; includes var. gracilis (Schleich. ex Roo k. & Grev.) R.R.Zander in VE, and var. icmadophila (Schimp. ex Mü1l.Ral.) R.R.Zander and var. rigidulus in EC D. taylori R.R.Zander; EC; 2400-3385 m D. umbrosus (Müll.Ral.) R.R.Zander [syn.: D. australasiae (Roo k. & Grev.) R.R.Zander var. umbrosus (Müii.Ral.) R.R.Zander]; VE; 2000-3300 m D. vinea/is (Brid.) R.R.Zander; VE; 1660-3600 m Erythrophyllastrum R.R.Zander E. andina (Sull.) R.R.Zander; CO, EC; 3600-4200 m Hennediella Paris H. bellii (E.B.Bartram) R.R.Zander; EC; 3110-3260 m H. denticulata (Wilson) R.R.Zander; EC; 3080-3090 m H. limbata (Mitt.) R.R.Zander; EC; 2200-4100 m Hymenostylium Brid. H. recurvirostrum (Redw.) Dixon; VE, CO; 950-4100 m Leptodontium (Müll.Ral.) Rampe ex Lindb.
Ref: Zander, 1972. L. araucarietii (Müli.Ral.) Paris; EC; 2100-4100 m L. brachyphyllum Broth. & Thér.; CO, VE, EC; 2650-4100 m L. capitu/igerum Müll.Ral.; CO, VE, EC, PE; 2200-4100 m L. erythroneuron Rerzog; CO, VE, EC; 3500-4050 m L.jlexifolium (Dicks. ex With.) Rampe; CR, CO, VE, EC;
2400-4525 m
[VOL. 84
L. luteum (Taylor) Mitt.; CO, VE, EC; 2420-4000 m L. planifolium Rerzog; CO, EC; 3300-3700 m L. proliferum Rerzog; CO; 3500 m , L. pungens (Mitt.) Kindb.; CO, VE, EC; 2265-5300 m L. stellaticuspis E.B.Bartram; CO; VE, EC; 3500-4470 m L. stoloniferum R.R.Zander; CR, CO, EC; 3450-3600 m L. subintegrifolium Thér. ex Rerzóg; EC; 4250 m L. syntrichioides (Müii.Ral.) Kindb.; CO, EC; 3080-3700 m L. viticulosoides (P.Beauv.) Wijk & Margad.; CR, CO, VE, EC; 1800-4000 m; includes var. exasperatum (Cardo!) R.R.Zander, var. sulphureum (Lorentz) R.R.Zander, and var. viticulosoides L. wallisii (Müli.Ral.) Kindb.; CR, CO, VE, EC; 3000-4270 m Mironia R.R.Zander [syn.: Morinia Cardo! (1910) nom. illeg., non A.Berl. & Bres. (1889)] M. ehrenbergiana (Müii.Ral.) R.R.Zander; CR, CO, VE, EC; 2700-4350 m; includes var. ehrenbergiana and var. elongata (Wi1son) R.R.Zander
.
Molendoa Lindb. M. andina (Mitt.) Broth.; EC; 3080 m Pseudocrossidium R.S.Williams P. excavatum (Mitt.) R.S.Williams; EC; 2700-3900 m P. replicatum (Tay1or) R.H.Zander; CO, VE, EC; 1030-4200 m Pseudosymblepharis Broth. P. schimperiana (París) R.A.Crum; CO, O:C; 1000-4000 m Sagenotortula R.R.Zander (syn.: Tortula p.p.) S. quitoensis (Taylor) R.R.Zander; CO, EC; 2500-4100 m Streptopogon Wilson S. calymperes Müli.Ral. ex Geh.; CO, VE, EC; 1000-4000 m S. erythrodontus (Spruce) Spruce ex Mitt.; CO, VE, EC; 1300-3900m Syntrichia Brid. (syn.: Tortu/a p.p.) S. aculeata (Wi1son) R.R.Zander; EC; 3810-4570 m S. amphidiacea (Müli.Ral.) R.R.Zander; VE, EC; 22003500m S. andicola (Mont.) Ochyra; CO, VE, EC; 3300-4340 m S. bogotensis (Rampe) Mitt.; CO, VE, EC; 2460-3950 m S.fragilis (Tay1or) Ochyra; CO, VE, EC; 2000-3690 m S. percarnosa (Müli.Ral.) Broth.; CO; 3520-4350 m S. pichinchensis (Taylor) R.R.Zander; CO, EC; 2700-4200 m S. princeps (DeNot.) Mitt.; EC; 4170 m S. ruralis (Redw.) F. Weber & Mohr; EC; 3700-4120 m Trichostomum Bruch T. aequatoriale Spruce ex Dixon; EC; 1830-4480 m T. brachydontium Bruch; CR, CO, VE, EC; 740-4000 m T. tenuirostris (Rook. & Taylor) Lindb.; CO, VE?, EC; 180-
3780m
1999]
PARAMOS
63
PRIONODONTACEAE
SPHAGNACEAE
Prionodon Müll.Hal. P. densus (Hedw.) Müll.Hal.; CO, VE, EC; 1450--4000 m P.fusco-lutescens Hampe; CO, VE, EC; 1750-3720 m
Sphagnum L. S. azuayense H.A.Crum; EC; 3350 m S. bals/evii H.A.Crum; EC; 2430-3630 m S. barclayae H.A.Crum; EC; 3750 m S. boyacanum H.A.Crum; CO; 3400 m S. cleefii H.A.Crum; CO; 3390 m S. compactum DC.; CO, VE; 1900-3600 m S. cundinamarcanum H.A.Crum; CO; 3300 m S. curvatulum H.A.Crum; CO, EC, 3050-3700 m S. cuspidatum Ehrh.; CR, CO, EC; 2500-3800 m S. cyclophyllum Su!!. & Lesq.; CO; 3375-3650 m S. incommodum H.A.Crum; EC; 2550-3450 m S. /aegaardii H.A.Crum; EC; 3900 m S. /ewisii H.A.Crum; CO; 3100 m S. limbatum Mitt.; CO, VE; 1680--4030 m S. maegdefraui H.Suzuki; VE; 3350 m S. magel/anicum Brid.; CR, CO, VE, EC; 1500-3980 m S. meridense (Hampe) Müll.Hal.; CO, VE, EC; 1220--3600 m S. oxyphyllum Wamst.; CO; 1400-3340 m S. pulchricoma Müll.Hal.; EC; 2800-3000 m S. pylaesii Brid.; EC; 3870 m S. recurvum P.Beauv. var. recurvum; CO; 1725-3800 m S. rio-negrense H.A.Crum; CO; 3350-3375 m S. sanctojosephense H.A.Crum & Crosby; CR, CO, VE, EC; 2200--4020 m S. sparsum Hampe; CR, CO, VE, EC; 1340-3800 m S. subsecundum Nees var. rufescens (Nees & Hornsch.) Hüb.; CR, CO, EC; 1400-3500 m S. sumapazense H.A.Crum; CO; 3500-3650 m S. tenerum Su11. & Lesq. ex Sull.; CO; 2460-3600 m
RACOPILACEAE Racopilum P.Beauv. R. intermedium Hampe; VE; 1800-3100 m R. tomentosum (Hedw.) Brid.; CR, CO; 245-3500 m
RHACOCARPACEAE Rhacocarpus Lindb. R. purpurascens (Brid.) Paris; CR, CO, VE, EC; 600--4200 m
RHIZOGONIACEAE Leptotheca Schwagr. L. boliviana Herzog; CO, EC; 1840-3700 m Pyrrhobryum Mitt. P. mnioides (Hook.) Manuel; CR, CO, VE; 2750-3670 m Rhizogonium Brid. R. novae-hollandiae (Brid.) Brid. [syn.: R. lindigii (Hampe) Mitt.]; CO, VE; 1500-3720 m
.RIGODIACEAE Rigodium Kunze ex Schwagr. R. toxarion (Schwagr.) A.Jaeger; CO, VE, EC; 2550--3750 m
SPLACHNACEAE
SELIGERIACEAE Blindia Bruch, Schimp. & W.Gümbel Ref.: Bartlett & Vitt, 1986. B. gradsteinii J.K.Bartlett & Vitt; CO; 4090 m B. magellanica Schimp.; CO, VE, EC; 2940--4200 m
Brachymitrion Taylor B.jamesonii Taylor; CO, VE, EC; 2770--4200 m B. moritzianum (Müll.Hal.) A.K.Kop.; CR, VE; 1780--3500 m Splachnum Hedw. S. weberbaueri Reim.; CO, VE, EC; 2500-3950 m
SEMATOPHYLLACEAE
Aptychella (Broth.) Herzog A. proligera (Broth.) Herzog; CO, VE, EC; 1800-3700 m
Tay/oria Hook. T. altorum Herzog; CO; 3400 m T. magellanica (Brid.) Mitt.; CO; 3200-3750 m T. rubricaulis A.K.Kop.; CO; 4260 m T. scabriseta (Hook.) Mitt.; CO, VE, EC; 2500--4340 m
Aptychopsis (Broth.) M.Fleisch. A. pycnodonta Herzog; CO; 3500 m
Tetraplodon Bruch & Schimp. T. mnioides (Hedw.) Bruch & Schimp.; CR, CO, VE, EC;
Acroporium Mitt. A. pungens (Hedw.) Broth.; CO; 80-3500 m
1800-3800 m Sematophyllum Mitt. S. aureo-sulfureum (Müli.Hal.) Broth.; CO; 2650-3690 m S. esmeraldicum (Müll.Hal.) Broth.; VE, EC; 2930-3700 m S. decumbens Mitt.; CO, EC; 1850-3800 m S. napoanum (DeNot.) Steere; CO, VE, EC; 1900--4000 m S. obliquerostratum Mitt.; EC; 3250 m S. swartzii (Schwagr.) Welch & H.A.Crum; VE; 2900--3300 m
THAMNOBRYACEAE Porotrichodendron M.F1eisch. P. nitidum (Hampe) Broth.; CO, EC; 2500-3700 m P. robustum Broth.; CO; 2340-3750 m P. superbum (Tay1or) Broth.; CO, VE, EC; 2400-3700 m
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MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
Porotrichum (Brid.) Hampe P. lancifrons (Hampe) I.Sastre; CO, VE; 1400-3250 m P. longirostre (Hook.) Mitt.; CO, VE; 500-3200 m
THUIDIACEAE Cyrto-hypnum (Hampe) Hampe & Lorentz C. leptocladium (Taylor) W.R.Buck & H.A.Crlim; EC; 1230-3690 m Thuidium Bruch & Schimp. T. delicatulum (Hedw.) Schimp.; CR, CO, VE, EC; 10003700 m T. peruvianum Mitt.; CO, VE, EC; 1840-4200 m T. phi/ibertii Limpr.; VE; 3000-3500 m T. pseudo-protensum (Müll.Hal.) Mitt.; CO, VE, EC; 22003000m
Acknowledgments Thanks are due to T. Pócs for severa! additional moss records for Venezuela and to Jesús Muñoz for data on Grimmia. A portion of this work was supported by grants from the National Science Foundation (DEB-88-18051, DEB-92-01281) to the first author, and the Andrew W. Mellon Foundation (grant to Dr. James L. Luteyn) funded a visit (for SPC) to the Smithsonian Institution.
Litera tu re Cited in Checklist of Mosses Balslev, H. & T. de Vries. 1991. Life forms and species richness in a bunch grass páramo on Mount Cotopaxi, Ecuador. Pp. 45-58. In: W. Erdelen, N. Ishwaran & P. Müller ( eds.), Tropical ecosystems. Margraf Scientific Books, Saarbrüken. Bartlett, J. K. & D. H. Vitt. 1986. A survey ofspecies in the genus B/india (Bryopsida, Seligeriaceae). New Zealand J. Bot. 24: 203-246. Bartram, E. B. 1931. A review ofthe American species of Da/tonia. Bull. Torrey Bot. Club 58: 31-48. - - - . 1953. Paramo mosses ofVenezuela and Colombia, collected by A. H. G. Alston. Bryologist 56: 165-168. - - - . 1964. Mosses ofCerro Antisana, Ecuadorian Andes. Rev. Bryol. Lichénol. 33: 1-14. Bowers, F. D. 1970. High elevation mosses of Costa Rica. J. Hattori Bot. Lab. 33: 7-35. Churchill, S. P. & E. L. Linares C. 1995. Prodromus bryologiae novo-granatensis. 2 vols. Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bibliot. José Jerónimo Triana 12: 1-924. ---,D. Griffin III & M. Lewis. 1995. Moss diversity of the tropical Andes. Pp. 335-346. In: S. P. Churchill, H. Balslev, E. Forero & J. L. Luteyn (eds.), Biodiversity
[VOL. 84
and conservation of neotropical montane forests. New York Botanical Garden, Bronx. Cleef, A. M. 1981. The vegetation of the páramos of the Colombian Cordillera Oriental.Diss. Bot. 41: 1-320. Fife, A. J. 1987. Taxonomic and nomenclatura! observations on the Funariaceae. 5. A revision of the Andean species of Entosthodon. Mem. NewYork Bot. Gard. 45: 301-325. Frahm, J.-P. 1989. The genus Chorisodontium (Musci, Dicranaceae) in the Neotropics. Trop. Bryol. 1: 11-23. ~~~. 1991. Dicranaceae: Campy1opodioideae, Paraleucobryoideae. Fl. Neotrop. Monogr. 54: 1-238. Fransén, S. 1995 [ 1996]. A taxonomic revision ofneotropical Bartramia section Vaginel/a C. Müll. Linbergia 20: 147-179. Griffin, D., III. 1979. Briófitos y líquenes de los páramos. . Pp. 79-87. In: Salgado-Labouriau (ed.), 1979a. --·- . 1986. Oreoweisia (Dicranaceae, Musci) in tropical ·: America: An annotated key to species. Cryptog. Bryol. Lichénol. 7: 433-438. -..- - . 1988. New World species of Breute!ia with erectappressed 1eafbases. Beih. Nova Hedwigia 90: 357-382. - - - . 1990a. F1oristics of the South American páramo moss flora. Trop. Bryol. 2: 127-132. - - - . 1990b. Two new pentastichous species of Zygodon from high e1evation in Venezuela. Cryptog. Bryol. Lichénol. 11: 163-168. Hyvi.inen, J. 1989. A synopsis of the genus Pogonatum (Polytrichaceae, Musci). Acta Bol. Fennica 138: 1-87. Lewinsky, J. 1984. Orthotrichum Hedw. in South America. l. Introduction and taxonomic revision oftaxa with immersed stomata. Lindbergia 10: 65-94. - - - . 1987. Orthotrichum (Orthotrichaceae) in South America. 2. Taxonomic revision of laxa with superficial stomata. Mem. New York Bot. Gard. 45: 326-370. Magdefrau, K 1983. The bryophyte vegetation ofthe forests and páramos of Venezuela and Colombia. Nova Hedwigia 38: 1-63. Malta, N. 1926. The genus Zygodon Hook. et Tayl. Acta Horti Bol. Univ. Latvian 1: 1-184. McFarland, K D. 1988. Revision of Brachythecium (Musci) for Mexico, Central America, South America, Antarctica, and Circum-subantarctic Islands. Ph.D. dissertation, University ofTennessee, Knoxville. Ochi, H. 1980. A revision of the neotropica1 Bryoideae, Musci (first part). J. Fac. Educ. Tottori Univ., Na!. Sci. 29: 49-154. - - - . 1981. A revision of the neotropical Bryoideae, Musci (second part). J. Fac. Educ. Tottori Univ., Nat. Sci. 30: 21-55. Ochyra, R. 1990. Gradsteinia andicola, a remarkable aquatic moss from South America. Trop. Bryol. 3: 19-28. Robinson, H. 1967. Pre1iminary studies on the bryophytes ofColombia. Bryologist 70: 1-43. Vareschi, V. 1970. Flora de los páramos. Talleres Gráficos Universitarios, Mérida. Zander, R. H. 1972. Revision of the genus Leptodontium (Musci) in the New World. Bryologist 75: 213-280. - - - . 1993. Genera of the Pottiaceae: Mosses of harsh environments. Bu!!. Buffalo Soc. Nat. Sci. 32: 1-378. - - - & A. M. Cleef. 1982. Studies on Colombian cryptogams XVI. Taxonomy and habitat of Kingiobryum paramico/a (Dicranaceae, Musci). Proc. Kon. Ned. Akad. Wetensch., Ser. C., 85: 627-634.
1999]
PARAMOS
65
Hepatics S. Rob Gradstein Richard Spruce, the great 19th-century explorer of the Andes and author of the classic Hepaticae Amazonícae et Andinae (Spruce, 1884--1885), was one ofthe first to collect hepatics in páramos. Surprisingly, he considered the zone quite poor in species (Spruce, 1886: 77): "La zone alpine des Andes est aussi pauvre en hépatiques que celle des Pyrenées." Spruce's negative impression ofhepatic diversity in the páramos was probably due to the fact that he visited only the rather dry, species-poor páramos of central Ecuador (Pichincha and Chimborazo provinces). His appreciation ofthe hepatic flora would undoubtedly have been different had he visited the species-rich, humid páramos of northern Ecuador, Colombia, or Venezuela. The most important hepatic collections made in páramos are those made by Antaine M. Cleef in Colombia during 1970-1972 (Gradstein, 1982, 1983, 1990; Gradstein et al., 1977; Gradstein & Hekking, 1979) and by Rudolf M. Schuster in Venezuela in 1976 (Schuster, 1978, 1978b, 1985, 1986, 1987, 1991, 1995). Other irnportant collectors include G. Wallis, H. Bischler, J. Cuatrecasas, J. Aguirre, K. Magdefrau, S. Winkler (Colombia), M. Allioni, E. Asplund, G. Harling, W. Jameson, H. Meyer (Ecuador), D. Griffin, J. Steyermark (Venezuela), G. Dauphin, N. Salazar Allen (Panama), A. M. Cleef et al., and G. Dauphin (Costa Rica). The author has collected hepatics in páramos in Costa Rica (years 1993, 1994), Colombia (years 1980, 1982, 1984, 1992), Ecuador (years 1976, 1988), and Peru (year 1982). At present 291 species of hepatics, in 88 genera and 34 families, are known from the neotropical páramos. Accepting about 1250 hepatic species in tropical America, in 189 genera (Salazar Allen et al., 1996), it appears that páramos harbar more than onefifth of the neotropical species and almost one-half ofthe genera. Taking into account the relatively small area occupied by the páramos, these figures indicate a very high diversity. Another striking feature ofthe hepatic flora ofthe páramos is the high degree of endemism. Seven genera and subgenera and about 70 species are exclusive!y known from páramos, accounting for 26.5% ofthe hepatic flora (Gradstein, 1998). The principal hepatic groups of the páramos are listed in Table IV. It appears that almost four-fifths of the neotropical hepatic families are represented in
the páramos; those lacking are usually small, specialized groups. Many families have only few taxa in páramos, however. It appears that the largest •mmber of species recorded per country by far have been those from Colombian páramos (230 spp.); the smallest nurnber are known from Panama (7 spp.). The relatively large area covered by páramo and the very moist climate may explain why Colombian páramos have so many species. On the other hand, Colombian páramos have also been studied more intensively than those of other countries. Therefore, the principal conclusion that can be drawn is that our knowledge of the páramo flora is very uneven and that much more collecting needs to be done to reach a reasonable leve! of understanding ofhepatic diversity in the páramos ofthe different countries. All taxa recorded at or abo ve 3200 m in the Andes and above 3000 m in Costa Rica have been included in the list ofspecies. Species occurring at lower elevations are excluded from this list unless it is stated that the record was from páramo. Elevational ranges of species are the total range ofthe taxon in the Neotropics. The information is based on the references cited at the end of the list and on herbarium specimens in SJU (records from Costa Rica), PMA (records from Panama), and U (all countries). The latter herbarium holds the largest collection ofpáramo hepatics worldwide and was a particularly rich source of information. In sorne instances it was uncertain whether the record was actually from páramo. This concerns particularly old references or old herbarium specimens, \acking details on locality or habitat. Additional fieldwork on the habitat preferences of the species is needed to help resolve these uncertainties.
CHECKLIST OF HEPATICS Anthocerotae ANTHOCEROTACEAE Phaeoceros Prosk. P. pichinchensis (Spruce) Haessel; EC; ?-3600 m
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MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
[VOL. 84
R. columbica (Steph.) Hassel ex Gradst.; CO; 1900-3700 m R. crassicaulis (Steph.) Meenks & DeJong; CO, PE; 19003600m Megaceros Campb. R.foliacea Meenks & DeJong; EC; 4000 m Megaceros sp.; CO; ?-4000 m R. hansmeyeri (Steph.) Meenks & DeJong; CO, VE, EC; 3200-4300 m R. herzogiana (Steph.) Meenks & DeJong; CO, VE, PE; 1400-4000 m Hepaticae R. judithae Meenks & DeJong; CO, PE; 3000-3400 m R. leptophylla (Spruce) Herzog; CO, VE, EC, PE; 1000ACROBOLBACEAE 3500m R. pallida (Spruce) Meenks & DeJong; CO, EC; 1500-4100 m Acrobolbus Nees R. papillata (Gottsche) Hasse1 ex Gradst.; CO, VE; 1950A. laceratus R.M.Schust.; VE; 3140 m 3450 m [Note: The record of R. insignis Schiffn. from Colombian páramo (Winkler, 1976) is erroneous and Lethocolea Mitt. refers to R. papillata (Meenks, 1987).] L. glossophylla (Spruce) Grolle; CR, CO, VE, EC, PE; 2400- , R. paramorum Meenks; CO; 3400-3800 m 3750 m R. parasitans (Steph.) Meenks & DeJong; CO, VE, EC, PE; 3000-4500 m Marsupidium Mitt. R. plumaeformis (Spruce) Méenks; CO, EC, PE; 1500-3700 m M. latifolium R.M.Schust.; VE; 3100 m R. poeppigiana (L.Lehm. & Lindenb.) Hassel ex Meenks & DeJong; CO, EC, PE; 2900?-4200 m Tylimanthus Mitt. R. smaragdina Meenks & DeJong; CO, VE; 2350-3600 m T. setaceo-ciliatus Steph.; CR?, CO, EC; 2800-4000 m R. trichomanoides (Spruce) Meenks; CO, EC, PE; 29003900 m ADELANTHACEAE R. wallisii (Steph.) Gradst.; CO; 3000?-4000 m
DENDROCEROTACEAE
Adelanthus Mitt. Ref.: Grolle, 1972; Schuster, 1978a, 1978b. A. aureomarginatus R.M.Schust.; CO, VE; 2000-3500 m A. crossii Spruce; CR, CO, VE; 3000-4250 m A. decipiens (Hook) Mitt. subsp. aureocinctus R.M.Schust.; VE; 3140 m [Note: Adelanthus crossii andA. decipiens subsp. aureocinctus may be untoothed high-altitude forros of A. decipiens (Hook) Mitt.] A. lindenbergianus (F.Lehm.) Mitt.; CR, PA, CO, VE, EC, PE; 1800-4200 m A. pittieri (Steph.) Grolle; CR, CO, VE; 2500-3500 m
ANEURACEAE Cryptothallus Malmb. Ref.: Crum & Bruce, 1997. C. hirsutus Crum; CR; 3100 m Riccardia Gray [Note: Riccardia is one of the most common hepatic genera in the neotropical páramos; the taxonomy and distribution ofthe species are still very incompletely known. The present list is largely based on Meenks, 1987. The lack ofrecords from Costa Rican páramo is an artifact; severa! (unidentified) species have been collected by the author and others.] Ref.: Meenks, 1987. R. aberrans (Steph.) Gradst.; CO, PE; 3000-3700 m R. algoides (Taylor) Meenks [syn.: R. squarrosa (Steph.) Gradst.]; CO, EC; 3000-3900 m R. capillacea (Steph.) Meenks & DeJong; CO, VE, EC?, PE; 2600-3900 m R. cilio/ata (Spruce) Gradst.; CO, VE, EC, PE; 2650-3750 m
ARNELLIACEAE Gongylanthus Lindb. G. granatensis (Gottsche) Steph.; CO, PE; 2500-4050 m G. liebmannianus (Lindenb. & Gottsche) Steph.; CR, PA, CO, VE, EC, PE; 2600-4300 m G. limbatus (Herzog) Grolle & Váiía (syn.: G. innovans S.Winkl., n.v.); CO, VE; 3100-4300 m
AYTONIACEAE Asterella P .Beauv. A. macropoda (Spruce) A.Evans; CR, CO, EC, PE; 23003700m
BALANTIOPSIDACEAE !sota chis Mitt. Ref.: Fulford, 1963. l. lacustris Herzog (syn.: Triandrophyllum maegdefraui S.Winkl., syn. nov.); CO, EC; 2500-4700 m l. lindigiana Gottsche; CO; 2300-3500 m l. lopezii (R.M.Schust.) Gradst. (syn.: Ruizanthus lopezii R.M.Schust.); CR, CO, VE, EC, PE; 3000-3800 m [Note: Ruizanthus lopezii ( fide Gradstein 974 7, Costa Rica, Cerro de la Muerte, on bank ofPanamerican Hwy. together with R. venezuelanus, 3000 m, 29 Sep 1995, GOET) has spiraled instead of straight capsule valves. The 2-lobed leaves of this species al so indicate a c1oser relationship to Isotachis than to Ruizanthus, which has 4-lobed 1eaves. Therefore, it is herewith transferred to Isotachis, as I.lopezii (R.M.Schust.) Gradst. comb. nov.
1999]
PARAMOS
(basionym: Ruizanthus lopezii R.M.Schust., Phytologia 39: 241. 1978).] l. multiceps Gottsche; CR, CO, VE, EC, PE; 1250-4100 m l. obtusa Steph.; VE, EC; 3200-3400 m [Note: Jsotachis lacustris and l. obtusa are characteristic species of shallow lakes and rivulets in neotropical páramos. As shown by Fulford (1963 ), the type specimens have very different leaf shapes (o vate in J. lacustris, orbicular in J. obtusa) and leaf incisions. However, additional collections made by the author and others show considerable variation and overlap in these characters, suggesting that the two may be conspecific. In the latter case, J. obtusa Steph. would be the older, hence corree! name.] l. serrulata (Sw.) Gottsche [syn.: l. haematodes (Lindenb. & Gottsche) Gottsche, l. madida (Hook. & Taylor) Mitt.]; PA, CO, VE, EC, PE; 700-4450 m [Note: Jsotachis haematodes (Lindenb. & Gottsche) Gottsche and J. madida (Hook. & Taylor) Mitt. are treated as synonyms of the common, highly variable l. serrulata following Gradstein et al., 1977.] Ruizanthus R.M.Schust. [Note: See note under Jsotachis lopezii.] R. venezuelanus R.M.Schust.; CR, CO, VE; 3000-4050 m
CAL YPOGEIACEAE Calypogeia Raddi Ref.: Fulford, 1968. C. andicola Bischler; CO, EC; 800-4100 m C. cyclostipa (Spruce) Steph.; CO, PE; 1700-3600 m C. peruviana Nees & Mont.; CO, EC, PE; 500-3800 m
CEPHALOZIACEAE Cephalozia (Dumort.) Dumort. Ref.: Váiía, 1988. C. bicuspidata (L.) Dumort.; CO; 3500-3800 m C. crossii Spruce (syn.: C. dussii Fulford); CR, CO, EC, PE; 500-4100 m C. pleniceps (Austin) Lindb.; CO; 3500-3700 m Odontoschisma (Dumort.) Dumort. O. atropurpureum Steph.; CO, VE, EC; 250?-4000 m O. denudatum (Nees) Dumort.; CO, EC; 600?-3500 m
CEPHALOZIELLACEAE Cephaloziella (Spruce) Schiffn. Ref.: Fulford, 1976; Schuster, 1978b. C. divaricata (Sm.) Schiffn. (syn.: C. andina Herzog); CO, VE, EC; 3400 m C.fragillima (Spruce) Fulford; CO, VE; 3000-3330 m C. granatensis (Jack) Fulford; CR, PA, CO; 2500-4000 m C. grisea R.M.Schust.; VE; 4150 m C. pungens Steph. ex Fulford; VE; 3600 m C. stolonifera R.M.Schust.; VE; 3650 m
67
FOSSOMBRONIACEAE Austrofossombronia R.M.Schust. Ref.: Schuster, 1994. A. peruviana (Gottsche) R.M.Schust. (syn.: Fossombronia ptychophyl/a Spruce, n.v.); CO, EC, PE; 3600-4000 m Fossombronia Raddi [Note: Neotropical Fossombronia has not been studied critically. The present list therefore like1y contains errors in identification.] F. brasi/iensis Steph.; CO, EC; 1800-4200 m F. paranapanemae Schiffn.; CO; 3000-4250 m F. sp.; CR; 3000 m
GEOCALYCACEAE Ref.: Fulford, 1976. Campanocolea R.M.Schust. Re f.: Schuster, 1997. C. fragmentissima (R.M.Schust.) R.M.Schust. (syn.: Lophocoleafragmentissima R.M.Schust.); CO, VE, EC, PE; 3200-4200 m Clasmatocolea Spruce Ref.: Engel, 1980. C. vermicularis (F.Lehm.) Grolle; CR, CO, VE, EC, PE; 1500-4500 m Heteroscyphus Schiffn. H. marginatus (Steph.) Fulford; CO, VE, EC; 1000-3300? m H. polyb/epharis (Spruce) Schiffn.; CO, VE, EC, PE; 23003500m Leptoscyphus Mitt. L. amphibolius (Nees) Grolle; PA; 1000-3150 m L. cleefii Fulford; CO; 3200-4100 m L. cuneifolius (Hook.) Mitt.; CR, PA, CO, VE, EC, PE; 2600-4400 m L.jackii (Steph.) Grolle; CO, EC; 3000-3900 m L. obcordatus (Spruce) Grolle; CO, EC; 3300-3500? m L. physocalyx (Rampe & Gottsche) Gottsche; CR, CO, VE, EC, PE; 2000-3850 m L.porphyrius(Nees) Grolle; PA, CO, VE, EC, PE; 100-4300m Lophocolea (Dumort.) Dumort. L. bidentata L. (syn.: L. coaduna/a (Sw.) Nees); PA, CO, VE, EC, PE; 100-3700 m L. erosa Gradst.; CR, EC; 3200-3700 m L. granatensis Gottsche; CO, PE; 1000-3700 m L. muricata (F.Lehm.) Nees; CO, VE; 1300-3400 m L. trapezoidea Mont.; CR, CO, EC, PE; 500-4000 m P/atycaulis R.M.Schust. Ref.: Schuster, 1995. P. renifolia R.M.Schust.; VE; ? m Pseudocephaloziella R.M.Schust. Ref.: Schuster, 1991. P. epiphytica R.M.Schust.; VE; 3140 m
68
MEMOIRS OF THE NEW YORK BOTAN! CAL OARDEN
GYMNOMITRIACEAE Gyinnomitrion Corda Ref.: Schuster, 1996a; Váiía, 1976. G. andinum (Herzog) Herzog; CO, PE; 3450-4200 m G. atrofi/um Váiía; CO; 3400-4000 m G. setacewn Grolle & Váiía; CR, CO; 3000-4300 m G. truncato-apicu/atum Herzog; CR, CO; 3400-4300 m Marsupel/a Dumort. Re f.: Schuster, I 996a. M. emarginata (Ehrh.) Dumort.; CO; 3150 m M. invo/uta Váiía; CO; 3600-4300 m M. microphyl/a R.M.Schust.; VE; 4160 m M. revo/uta (Nees) Dumort.; VE; 4400-4500 m M. tro/lii Herzog; CR, CO; 3500-4400 m M. xenophyl/a R.M.Schust. [syn.: Nanomarsupella xenophyl/a (R.M.Schust.) R.M.Schust.]; VE; 4160 m Paramomitrion R.M.Schust. Ref.: Schuster, l996a. P. paradoxum R.M.Schust.; VE; 4160 m Stephaniel/a Jack Ref.: Schmitt & Wink1er, 1969. S. paraphyllina Jack; CR, CO, VE, EC, PE; 2700-4500 m S. rostrata U.Schmitt; CR, CO; 3000-4300 m Stephanie/lidium S.Winkl. ex Grolle S. sleumeri (Mü1l.Frib.) S.Winkl. ex Grolle; CO; 3100-? m
HAPLOMITRIACEAE Haplomitrium Nees Ref.: Bartho1omew-Began, 1992. H. blumei Nees [syn.: H. andinum (Spruce) R.M.Schust.]; EC; 3900 m
I[VOL. 84
lombia, seems to be a synonym of Jsotachis lacustris judging from the original description and illustration.] T. subtrifidum (Hook. & Taylor) Fulford & Hatcher; CR, PA, CO, VE, EC, PE; 1800-4500 m
JUBULACEAE Frullania Raddi Ref.: Stot1er, 1969; Yuzawa, 1991. F. a/bertii Steph.; CO, EC; 3400-3950 m [Note: F. albertii is probably a robust, high-altitude form ofF. standaertii Steph. Both taxa are characterized by the rather large, lanceolate stylus (Yuzawa, 1991).] F. 1,1recae (Spreng.) Gottsche; CO, EC; 800-3800 m [Note: The páramo records ofF. a recae are in need ofrevision. As shown by Yuzawa (1991), the species has been confused with both F. ecklonii (Spreng.) Spreng. and F. viminicola Spruce.] F. brasiliensis Raddi; CO, EC; 200-3800 m F. convoluta Lindenb. & Hampe; P A, CO, VE, EC, PE; 1900-4000 m F. cuencensis Taylor; CO, EC; 2100-3400 m F. dusenii Steph.; CO, EC; 1600-3400 m F. holostipula Hatt. & D.G.Griffin; VE; 2600-3300 m F. lobato-hasta/a Steph.; CO; 1800-3500 m F. peruviana Gottsche; CO, VE, EC, PE; 1500-3700 m [Note: Two species of Frullania sect. Meteoriopsis are recognized in neotropical páramos: F. convoluta (with rounded to obtuse 1eafapex) andF. peruviana (with acule to short acuminate leaf apex). Both species have deep1y cordate 1eafbases with two large auricles.] F. planifolia Steph.; CR; 3000 m F. pluricarinata Gottsche; CO, EC; 1800-3350 m F. sphaerocephala Spruce; CR, CO, EC, PE; 2700-4000 m F tetraptera Nees & Mont.; CO, VE, EC, PE; 3100-3850 m
JUNGERMANNIACEAE HERBERTACEAE Herbertus S.Gray Ref.: van Reenen, 1982. H. acanthe/ius Spruce (syn.: H. limbatus Steph.); CR, CO, EC, PE; 2400-4350 m H. co/ombianus Reenen; CO; 3250-3750 m H.juniperoides (Sw.) Grolle; CR, PA, CO, EC, PE; 20003600m H. oblongifolius (Steph.) Gradst. & C1eef; CO; 4060 m H. subdentatus (Steph.) Fu1ford (syn.: H. subnivalis S.Winkl.); CR, PA, CO, VE, EC, PE; 2400-4350 m 0/gantha R.M.Schust. Ref.: Schuster, l996b. O. eophyl/a R.M.Schust.; EC; 4100 m Triandrophyllum Fu1ford & Hatcher [Note: Triandrophyllum maegdefraui S.Winkl., described from the Sierra Nevada de Santa Marta, Co-
Anastrophyllum (Spruce) Steph. Ref.: Váiía, 1980, 1984; Gradstein & Váiía, 1987. A. auritum (F.Lehm.) Steph.; CR, CO, VE, EC, PE; 20004500 m A. austroamericanum Váiía (syn.: Marsupe/la austroamericana Váiía, nom. inval.); CO, VE; 3750-4300 m A. leucocepha!um (Tay1or) Steph.; CO, EC, PE; 2300-4250 m A. minutum (Schreb.) R.M.Schust. [syn.: A. gemmiferum S.Winkl., syn. nov.]; CO, VE; 3700-4500 m [Note: Anastrophyllum gemmiferum S.Winkl. (Type: Colombia, Sierra Nevada de Santa Marta, leg. S. Winkler, ULM) is treated as a synomym of A. minutunz (Schreb.) R.M.Schust., at the advice of J. Váiía (in 1itt.).] A. nigrescens (Mitt.) Steph.; CR, CO, VE, EC, PE; 20004300m A. pearcei (Steph.) R.M.Schust.; CO, VE, PE; '?m A. stellatunz R.M.Schust. (syn.: A. gradsteinii Váiía, nom. inval.); CR, CO, VE, EC, PE; 3140-4000 m A. tubu/osum (Nees) Grolle; CO, VE, EC, PE; ? m
1999]
69
PARAMOS
Andrewsianthus R.M.Schust. A. jamesonii (Mont.) Váiía [syn.: A. kilimajaricus (S.W.Arnell) Grolle & Váiía; Lophozia incisa (Schrad.) Dumort. subsp. austrigena R.M.Schust., n.v.); CO, VE, EC, PE"; 2000-4750 m Cryptochila R.M.Schust. Ref.: Grolle, 1971. C. grandiflora (Lindenb. & Gottsche) Grolle; PA, CO, VE, EC, PE; 1300-4630 m Gymnocoleopsis (R.M.Schust.) R.M.Schust. G. multiflora (Steph.) R.M.Schust.; CO, VE, PE; 3500-4400 m
S. liberata Inoue; CR, CO; 3000-3900 m S. manca (Mont.) Steph.; CO, EC, PE; 3000?-3500 m
LEJEUNEACEAE Amphilejeunea R.M.Schust. Re f.: Schuster, 1986. A. patellifera (Spruce) R.M.Schust.; CR; 2500-3200 m A. viridissima R.M.Schust.; CO, VE; 3000-4150 m Anoplolejeunea (Spruce) Schiffn. A. conferta (Meissn.) A.Evans; CR, CO, VE, EC, PE; 6003700 m
Jamesoniella (Spruce) Carring Ref.: Grolle, 1971. J. autumnalis (DC.) Steph.; CO, VE; 2000-3500 m J. rubricaulis (Nees) Grolle; CR, PA, CO, VE, EC, PE; 1350-4300 m J. undata (Mont.) Steph.; CO, PE; 1900-4300 m
Aureolejeunea R.M.Schust. Ref.: Schuster, 1986, 1987. A. aurifera R.M.Schust.; CO, VE; 3500-3700 m A. paramicola (Herzog) R.M.Schust. [syn.: Aureolejeunea paramoensis R.M.Schust.; Omphalanthus paramicola (Herzog) Gradst.]; CO, VE, EC; 2000-3700 m A. quinquecarinata R.M.Schust.; CO, VE; 3000-4000 m
Jungermannia L. Ref.: Váiía, 1973, 1974. J. decolor Schiffn.; CO, PE; 500-3500 m J. hyalina Lyell; CO; 3150 m J. linguifolia Gottsche; CO, EC, PE; 2500-3600 m J. ovato-trigona (Steph.) Grolle; CO, EC, PE; 2800-4200 m J. sphaerocarpa Hook.; CR, CO, VE, EC, PE; 2000-4100 m
Blepharolejeunea R.M.Schust. Ref.: Gradstein, 1994. B. incongrua (Lindenb. & Gottsche) Van S1ageren & Kruijt; CR, CO, VE, EC, PE; 1800-4100 m B. securifolia (Spruce) R.M.Schust.; CR, CO, VE, EC, PE; 3000-4600 m
Lophonardia R.M.Schust. [Note: According to J. Váiía (in litt.), Lophonardia is a doubtfu1 taxon, probab1y not meriting recognition as a separate genus.] L. caespitosa R.M.Schust.; VE; 4150 m Lophozia (Dumort.) Dumort. L. incisa (Schrad.) Dumort.; CR, CO, VE, EC, PE; 25004500m L. laxifolia (Mont.) Grolle (syn.: L. subinflata Steph.); CR, CO, VE, EC; 3000-4100 m L. stolonifera R.M.Schust.; VE; ? m L. verruculosa R.M.Schust.; VE; ca. 3150 m Nardia Gray Ref.: Enge1, 1988. N succulenta (Rich. ex F.Lehm.) Spruce; CO, EC; 800--3300 m
Brachiolejeunea (Spruce) Schiffn. Ref.: Van Slageren, 1985; Gradstein, 1994 B. laxifolia (Taylor) Schiff¡¡.; CR, CO, VE, EC, PE; 15003800 m Cheilolejeunea (Spruce) Schiffn. [Note: Two additional species of Cheilolejeunea from the páramos ofCo1ombia were described by Schuster ( 1992). Since Latin descriptions were not provided, the names are invalid and have been omitted from the present list.] C. choachina (Gottsche) Gradst., comb. nov. [syn.: Lejeunea choachina Gottsche, Ann. Sci. Na!. Bot., Sér. 5, 1: 156. 1864; Strepsilejeunea choachina (Gottsche) Steph.]; CO; 3000-4300 m C. erostrata R.M.Schust.; VE; 3140 m C. laevicalyx (Jack & Steph.) Grolle; CO; 3300 m
Rhodoplagiochila R.M.Schust. [Note: Following Inoue (1984), Rhodoplagiochila is p1aced in the Jungermanniaceae instead of P1agiochi1aceae.] Ref.: Schuster, 1978a; lnoue, 1984. R. rosea R.M.Schust.; VE; 3700-3750 m
Calura Nees C. calyptrifolia (Hook.) Dumort.; VE; 3200+ m C. naumannii Steph. (syn.: C. patagonica Ast); CO, EC; 3200-3800 m C. ornithocephala Herzog; CO, VE, EC; 3300-3800 m C. tenuicornis (A.Evans) Steph.; VE; 2300-3450 m
Syzygiella Spruce Ref.: Inoue, 1966. S. anomala (Lindenb. & Gottsche) Steph.; CR, CO, EC, PE; 2000-3600 m S. campanulata Herzog; CR, CO, VE; 2000-3750 m S. integerrima Steph.; CO; 1000-3550 m
Diplasiolejeunea (Spruce) Schiffn. D. alafa Ast.; VE; 2900-3300 m D. involuta S.Winkl. subsp. andicola Pócs; VE; 3300 m D. papilionacea R.M.Schust.; VE; 2900-3450 m D. pauckertii (Nees) Steph.; CO, VE, PE; 2900-3340 m
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MEMOIRS OF THE NEW YORK BOTAN! CAL GARDEN
Drepanolejeunea (Spruce) Schiffn. Ref.: Bischler, 1964. D, andina Herzog; CO; 3500-4100 m D. araucariae Steph.; VE; 3050-4000 m D. aurita Bischler; EC; 3350-3450 m D. granatensis (Jack & Steph.) Bischler; CO, VE; 20003700 m . D. navicularis Steph.; CO, VE; 3000-3700 m Frullanoides Raddi Ref.: Van Slageren, 1985; Gradstein, 1994. F. densifolia Raddi; CO, EC, PE; 500-3650 m Harpalejeunea (Spruce) Schiffn. ?H. ancistrodes (Spruce) Schiffn.; CO, EC; 3000-3800 m. H. cinchonae (Nees) Schiffn.; CO, VE, PE; 700-3400 m H. grandis Grolle; CO; 3750 m H. verrucosa Herzog; VE; 3200 m H. sp. nov.; CO; 3200-3680 m Leucolejeunea A.Evans L. xanthocarpa (F.Lehm. & Lindenb.) A.Evans; CO, VE, EC; 500-3500 m Lindigianthus Kruijt & Gradst. Ref.: Gradstein, 1994. L. cipaconeus (Gottsche) Kruijt & Gradst. [syn.: Dicranolejeunea cipaconea (Gottsche) Steph.]; CR, CO, VE, EC, PE; 1700-3500 m Macrolejeunea (Spruce) Schiffn.
M. pallescens (Mitt.) Schiffn.; CR, CO, VE, EC, PE; 20003800m Microlejeunea (Steph.) Schiffn.
M. bullata (Tay1or) Steph.; CR, CO, VE, EC; (0-)1000-3600 m M. colombiana Bischl.; VE; 3090-4070 m Omphalanthus Nees O.filiformis (Sw.) Nees; CR, CO, EC, PE; 1200-3900 m O. platycoleus Herzog; CO; 3000?-3750 m
LEPICOLEACEAE Lepicolea Dumort. Ref.: Fulford, 1963. L. ochroleuca (Spreng.) Spruce; CR; 1600-3100 m L. pruinosa (Taylor) Spruce; CR, CO, VE, EC, PE; 17003800m L. ramentifissa Herzog; VE; 2900-3300 m
LEPIDOZIACEAE Bazzania S.Gray [Note: The neotropical species of Bazzania and Lepidozia are poorly known and in need ofrevision; the present
[VOL. 84
list like1y contains misidentifications, and sorne species names may prove to be synonyms.] Ref.: Fulford, 1963. B. arcuata (Lindenb. & Gottsche) Trevis.; CR, CO, EC; 2003600m B. canelensis (Steph.) Fulford; CO; ?-3750 m B. chilensis (Steph.) Fulford; CO, EC; 2500-3600 m B. crassidentata Fulford; CO; 3500-3600 m B. diversicuspis Spruce; CO; 100-3500 m B. falca/a (Lindenb.) Trevis.; CO; 1000-3600 m B. hookeri (Lindenb.) Trevis. (syn.: B. robusta Spruce); CO, VE, PE; 100-3700 m B. jamaicensis (F.Lehm. & Lindenb.) Trevis.; CR, CO; 1500-3500 m B. latidens (Gottsche) Fu1ford; CO; 3750 m B . .longistipula (Lindenb.) Trevis.; CO, EC; 750-3500 m B. placophylla (Taylor) Grolle; CO, EC, PE; ?-3750 m B. roraimensis (Steph.) Fu1ford; CO, PE; 1500-3600 m B. taleana (Gottsche) Fulford; VE; 3200+ m Kurzia G.Martens Ref.: Pรณcs, 1984. K. capillaris (Sw.) Grolle (syn.: Kurzia verrucosa Steph.); CR, CO, VE, EC, PE; 500-3800 m K.jlagellifera (Steph.) Grolle; CO; 1500-3700 m Lepidozia (Dumort.) Dumort. [Note: The neotropical species of Bazzania and Lepidozia are poorly known and in need ofrevision; the present list 1ike1y contains misidentifications, and sorne species names may pro veto be synonyms.] Ref.: Fulford, 1966. L. alstoni Fulford; CO, EC; 3200-4500 m L. andicola Beauverd; CO, VE, EC; 2650-3600 m L. auriculata Mitt. ex Steph.; CO, EC, PE; 3300?-4000 m L. caespitosa Spruce; CO, VE, EC, PE; 1200-3800 m L. cupressina (Sw.) Lindenb.; PA, CO, VE, PE; 1000-3600 m L. macrocolea Spruce; CO, VE, EC, PE; 1000-4000 m L. peruviensis Steph.; CO, VE, EC, PE; 2500-3650 m L. wal!isiana Steph.; CO, VE, EC; 1000-3400 m Paracromastigum Fulford & J.Tayl. P. bifidum (Steph.) R.M.Schust.; VE; 2000-3400 m P. granatense (Gottsche) R.M.Schust. [syn.: Bonneria granatensis (Gottsche) Fulford & J.Tayl.]; CR, CO; ?-3200 m Pseudocephalozia R.M.Schust. Ref.: Schuster & Engel, 1974. P. quadriloba (Steph.) R.M.Schust.; CR, CO, EC, PE; 32004100 m Telaranea Spruce ex Schiffn. Re f.: Schuster, 1978a. T. microstipulata R.IVLSchust.; VE; ? m T. nematodes (Gottsche ex Austin) Howe; CR, PA, CO, VE, EC, PE; 200-4250 m T. quadrifida R.M.Schust.; VE; '! m
1999]
PARAMOS
71
brogniartii Mont.; CO, VE, EC; 700-3600 m marginata Steph.; CO; 800-4000 m podophylla (Thunb.) Mont. & Nees; CO; 1800-3850 m rubescens Steph.; CO; ?-3600 m
Dumortiera Nees D. hirsuta L.; CO, EC; 150-3500 m
S. S. S. S.
Marchan tia L. Ref.: Bisch1er, 1984. M. berteroana F.Lehm. & Lindenb.; CR, CO; 1800-3600 m M. plicata Nees & Mont.; CR, CO, EC, PE; 1700-3800 m M. polymorpha L.; CO; 3000-3800 m
Noteroclada Tay1or ex Hook. & Tay1or N. conjluens Taylor ex Hook. & Wilson; CR, CO, VE, EC, PE; 1800-4000 m
MARCHANTIACEAE
PELLIACEAE
METZGERIACEAE
PLAGIOCHILACEAE
Metzgeria Raddi [Note: The neotropica1 species of Metzgeria are still poor1y known, notwithstanding the work by Kuwahara (1986); the present 1ist shou1d be considered pre1iminary.] Ref.: Kuwahara, 1986. M. albinea Spruce; CO, EC; 500-3800 m M. atramentaria Kuwah.; CO; 3450--4000 m M. attenuata Steph.; CO; 3350-3600 m M. bischlerae Kuwah.; CO; 3500-3600 m M. cleefii Kuwah.; CO; 3300 m M. consanguÃnea Schiffn.; CO, VE, PE; 2100-3500 m M. decipiens (C.Massal.) Schiffn.; CO, PE; 500-3900 m M. dorsipara (Herzog) Kuwah.; CO, EC; 3350-3900 m M.filicina Mitt.; CO, EC, PE; 2000-3300 m M. gigantea Steph.; CR, CO, PE; 1500--4350 m M. lechleri Steph.; CO, PE; 3000-3800 m M. leptoneura Spruce; CO, VE, EC, PE; 500--4200 m M. liebmanniana Lindenb. & Gottsche; CO, VE, EC, PE; 1500-4000 m M. metaensis Kuwah.; CO; 4000-4200 m M. neotropica Kuwah.; CO, EC, PE; 3100--4000 m M. papulosa Steph.; CO, VE; 700-3600 m M. parviinvolucrata Kuwah.; PE; 3400 m M. polytricha Spruce; CO, PE; 1000--4000 m M. procera Mitt.; CO; 200-3600 m M. sandei Schiffn.; CO; 650-3950 m
Plagiochila (Dumort.) Dumort. [Note: The neotropical species of Plagiochila are very poorly known. The present list is large1y based on identifications of the late Dr. H. Inoue, whose revision ofthe group remained unfinished at the time of his death in 1989.] Ref.: Inoue, 1987, 1989; Robinson, 1967. P. cleefii Inoue; CO; 4050 m P. cuatrecasii H.Rob.; CO; 3600-4200 m P. dependula Taylor [syn.: Jamesoniella dependula (Taylor) Steph.]; CO, EC; 3500-4200 m P. echinella Gottsche; CO, VE?, EC, PE; 2400-3750 m P. fuscolutea Taylor (syn.: P. jelskii Loitl., P. scopulosa Steph.); CR, CO, VE, EC, PE; 2600-4000 m P. guevarii H.Rob. (syn.: P. dana-griffinii Inoue, n.v.); CO, VE; 3300-3700 m P. jamesonii Taylor; CO, EC; 2000-3600 m P. jarami/loi H.Rob.; CR, CO; 3000-3520 m P. /ongispina Lindenb. & Gottsche; CR, CO, VE, EC; 28003700m P. oblita Steph.; CO; 3750 m P. ovala Lindenb. & Gottsche; CR, CO; 2500-3750 m P. pachyloma Taylor; CO, EC, PE; 2000-3600 m P. paraphy/lina Herzog; CO; 3200 m P. revo/vens Mitt.; CO, EC; 4000-4300 m P. stolonifera Lindenb. & Gottsche; CR, CO; 1900-4200 m P. triangulifolia Steph.; CO; 4050 m P. umbrosa Steph.; CO; 3000-3750 m P. verrucu/osa R.M.Schust.; CO; 1500-3300 m
MONOCLEACEAE Monoclea Hook. Ref.: Gradstein et al., 1993. M. gottschei Lindb. subsp. elongata Gradst. & Mues; CR, CO, EC; 150-3500 m
PALLAVICINIACEAE Jensenia Lindb. Ref.: van der Gronde, 1980. J. erythropus (Gottsche) Grolle; CR, PA, CO, VE, EC, PE; 2200-4100 m J. jlorschuetzii van der Gronde; CO, EC; 4000-4100 m Symphyogyna Nees & Mont. Ref.: Uribe & Aguirre, 1995. S. brasiliensis Nees & Mont.; CR, CO, EC; 1000-3300 m
Steereochila Inoue Ref.: Inoue, 1988; Gradstein et al., 1994. S. ecuadorica Inoue; CR, EC; 2500?-3200 m
PLEUROZIACEAE Pleurozia Dumort. Ref.: Thiers, 1993. P. paradoxa Jack [syn.: Eop/eurozia paradoxa (Jack) R.M.Schust.]; CO, VE, EC; 2500-3800 m
PORELLACEAE Parella L. Ref.: Swai1s, 1970.
MEMOIRS OF THE NEW YORK BOT ANICAL GARDEN
72
P. leiboldii (F.I.,ehm. & Lindenb.) Trevis.; CR; 1360-4000 m P. squamulifera (Tay1or) Trevis.; CO, VE, EC, PE; 22003900 m
PSEUDOLEPICOLEACEAE
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T.fi/icau/is Steph.; CO; 800-3500 m T. robusta Steph.; CO; 2800-360 m T. sprucei Steph.; CO, PE; 1800-3600 m T. tomentosa (Sw.) Gottsche; CR, PA, CO, VE, EC, PE; 1003600m
Ref.: Schuster, 1985.
B/eph'arostoma (Dumort.) Du~ort. B. trichophy/lum (L.) Dumort.; CR, CO, VE, PE; 2700-4000 m Temnoma Mitt. T. chaetophyl/a R.M.Schust.; CO, VE; 3000-3700 m
RADULACEAE Radula Dumort. Ref.: Jans, 1980; Yamada, 1988; Reiner-Drehwald, 1994. R. episcia Spruce (syn.: R. cornucopiae Spruce); CO, EC; 1000-3600 m R.frondescens Steph.; CO, VE; ?-3800 m R.jamesonii Taylor; CO, EC; ?-3700 m R. nudicaulis Steph.; CO; 1500-4000 m R. sonsonensis Taylor; CO; 3000-4200 m R. voluta Tay1or (syn.: R. ramulina Taylor); CR, CO, EC, PE; 1500-4200 m
RICCIACEAE Riccia L. R. lame/losa Raddi (syn.: R. indusiata S.Winkl., syn. nov.); CO; 4200 m [Note: Riccia indusiata S.Winkl., described from the Sierra Nevada de Santa Marta, Colombia, is a synonym of R. lame/losa Raddi (fide holotype in ULM; identity confirmed by S. Jovet-Ast, Paris).] R. sorocarpa Bisch.; PE; 1500-4550 m
SCAP ANIACEAE Ref.: Gradstein & Váña, 1987.
Dip/ophyllum (Dumort.) Dumort. D. andico/um R.M.Schust.; VE; 4200 m D. obtusatum (R.M.Schust.) R.M.Schust.; CO, VE, EC, PE; 2500-4200 m Scapania (Dumort.) Dumort. S. cuspiduligera (Nees) Müll.Frib.; CO; 4300 m S. portoricensis Hampe & Gottsche; CR, CO, VE, EC, PE; 1500-3750 m
TRICHOCOLEACEAE Trichocolea Dumort. [Note: The taxonomy ofneotropical Trichocolea is still incompletely known; the present lis! like1y contains misidentifications, and sorne species names may prove to be synonyms. Most species listed are forest laxa occurring only accidentally in páramo.] Ref.: Fulford, 1963. T. elliotii Steph.; CO; l 000-3650 m
Acknowledgments This paperwas drafted while the author was an Andrew W. Mellon Visiting Scientist at the Smithsonian Institution, Washington. I am grateful to the Andrew W. Mellon Foundation for making m y visit possible (grant to James L. Luteyn) and to the Director and staff of the Botany Department, Smithsonian Institution, for support and use of their facilities. Thanks are also dueto Gregorio Dauphin (SanJosé) forproviding additional records from Costa Rica and Panama, to Tamás Pócs (Eger) for additional records from Venezuela, to Andrea Lücking (Ulm) for loan of the type of Riccia indusiata, and to Antoine Cleef, Jim Luteyn, and Orlando Rangel for valuable suggestions.
Literature Cited in Checklist of Hepatics Bartholomew-Began, B. 1992. A morphogenetic re-evaluation of Haplomitrium Nees (Hepatophyta). Bryophyt. Biblioth. 41: l-297. Bischler, H. 1964. Le genre Drepano/ejeunea Stephani ~n Amérique Centrale et Méridionale. Rev. Bryol. Lichénol. 33:15-179. - - - . 1984. Marchantia L. The New World species. Bryophyt. Biblioth. 26: 1-228. Crum, H. & J. Bruce. 1997. A new species ofC!yptothallus from Costa Rica. Bryologist 99: 433-438. Engel, J. J. 1980. A monograph ofthe genus Clasmatocolea. Fieldiana, Bot. n.s. 3: 1-229. - - - . 1988. The taxonomic position of Apotomanthus (Hepaticae). Beih. Nova Hedwigia 90: 203-221. Fulford, M. H. 1963. Manual ofthe leafy Hepaticae ofLatin America. I. Mem. New York Bot. Gard. 11: 1-172. - - - . 1966. Manual of the leafy Hepaticae of Latin America. II. Mem. New York Bot. Gard. 11: 173-276. - - - . 1968. Manual of the leafy Hepaticae of Latin America. III. Mem. New York Bot. Gard. 11: 277-392. ----. 1976. Manual of the leafy Hepaticae of Latin America. IV. Mem. New York Bot. Gard. 11: 393-535. Gradstein, S. R. (ed.). 1982. Studies on Colombian Cryptogams I-X. Institute ofSystematic Botan y, University of Utrecht. - - - . 1983. Studies on Colombian Cryptogams XI-XX. Institute ofSystematic Botany, University ofUtrecht. - - - (ed.). 1990. Studies on Colombian Cryptogams XXIXL. Institute ofSystematic Botan y, University ofUtrecht. - - - . 1994. Lejeuneaceae: Ptychantheae, Brachiolejeuneae. Fl. Neotrop. Monogr. 62: 1-225. - - - . 1998. Hepatic diversity ofthe neotropical páramos. Monogr. Syst. Bot. Missom:,i Bot. Gard. 68: 69-85. - - - & W. H. A. Hekking. 1979. A catalogue of the Hepaticae ofColombia. J. Hattori Bot. Lab. 45: 93-144. - - - & J. Váiía. 1987. On the occurrence of Laurasian
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liverworts in the tropics. Mem. New York Bot. Gard. 45: 388--425. ---,M. H. Fulford & A. M. Cleef. 1977. Oil body structure and ecological distribution of selected species of tropical Andean Jungermanniales. Proc. Kon. Ned. Acad. Wetensch. ser. C, 80: 377--420. ---,R. Klein, L. Kraut, R. Mues, J. Sporle & H. Becker. 1993. Phytochemical and morphological support for the existence of two species in Monoc/ea (Hepaticae ). PI. Syst. Evol. 180: 115-135. - - - , A Lücking, M. Morales & G. Dauphin. 1994. Additions to the hepatic flora ofCosta Rica. Lindbergia 19: 73-86. Grolle, R. 1971. Jamesonie/la und Verwandte. Feddes Rep. 82: 1-99. - - - . 1972. Zur Kenntnis von Ade/anthus Mitt. J. Hattori Bol. Lab. 35: 325-370. Inoue, H. 1966. A monograph ofthe genus Syzygie/la Spruce. J. Hattori Bot. Lab. 29: 171-213. - - - . 1984. The genus Plagiochila (Dum.) Dum. in Southeast Asia. Akademia Scientific Book, Tokyo. - - - . 1987. New species of Plagiochila (Dum.) Dum. from the Neotropics. Pp. 95-105. In: H. Inoue (ed.), Studies on cryptogams in southem Peru. Tokai University Press, Tokyo. - - - . 1988. Steereochila, a new genus ofthe Plagiochilaceae from the Neotropics. Mem. New York Bot. Gard. 45: 279-282. - - - . 1989. Studies on some Plagiochila species in the Neotropics. Bull. Nat. Sci. Mus., Tokyo, Ser. B, 15:35--47. Jans, E. 1980. High Andean species of Radula (Hepaticae). Proc. Kon. Ned. Acad. Wetensch. ser. C, 82:421--432. Kuwahara, Y. 1986. The Metzgeriaceae ofthe Neotropics. Bryophyt. Biblioth. 28: 1-254. Meenks, J. J. D. 1987. A guide to the tropical Andean species of Riccardia (Hepaticae). J. Hattori Bot. Lab. 62: 161-182. Pócs, T. 1984. Synopsis of the African Lepidoziaceae. Proc. Third Meeting Bryol. Centr. East. Europe, Praha: l 07-119. Reiner-Drehwald, E. 1994. El género Radula Dum. (Radulaceae, Hepaticae) en el noreste de Argentina. Trop. Bryol. 9: 5-22. Robinson, H. 1967. Preliminary studies on the bryophytes of Colombia. Bryologist 70: 1--61. Salazar Al!en, N., S. R. Gradstein & S. P. Churchill. 1996. A guide to the bryophytes of Tropical America: A report. Anales Inst. Biol. Univ. Nac. Autón. Mexico, Ser. Bot. 67: 59-65. Schmitt, U. & S. Winkler. 1969. Systematische Untersuchungen über die foliase Lebermoosgattung Stephaniel/a Jack. Osterr. Bot. Zeitschr. 115: 120-135. Schuster, R. M. 1978a. Studies on V enezuelan Hepaticae. l. Phytologia 39: 239-251. - - - . 1978b. Studies on Venezuelan Hepaticae. II. Phytologia 39: 425--432. - - - . 1985. Studies on Venezuelan Hepaticae. III. FamiIies Blepharostomataceae and Ba1antiopsidaceae. Nova Hedwigia 42: 49-79. - - - . 1986. Studies on Venezuelan Hepaticae. IV. On Amphilejeunea Schust. and Aureo/ejeunea Schust. Nova Hedwigia 44: 1-23. - - - . 1987. On Aureo/ejeunea Schust. and Brachiolejeunea paramicola Herzog. Phytologia 61: 445--44 7. - - - . 1991. Studies on Venezue1an Hepaticae. V. On Pseudocephaloziella epiphytica Schust. Nova Hedwigia 53: 331-339. - - - . 1992. The oi1-bodies ofthe Hepaticae. Il. Lejeuneaceae (Part 2). J. Hattori Bot. Lab. 72: 163-359.
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- - - . 1994. Studies on Metzgeria1es. III. The c1assification of the Fo·ssombroniaceae and on Austrofossombronia Schust., gen. n. Hikobia 11: 439--449. - - - . 1995. Venezuelan Hepatics. VI. On P/atycaulis Schust. (Jungermanniales). Nova Hedwigia61: 391-396. - - - . l996a. Studies on antipodal Hepaticae. XII. Gymnomitriaceae. J. Hattori Bot. Lab. 80: 1-147. - - - . 1996b. On Olgantha Schust., gen.n. Nova Hedwigia 63: 529-543. - - - . 1997. On Campanocolea Schust. J. Hattori Bot. Lab. 82: 253-259. - - - & J. J. En gel. 1974. A monograph ofthe genus Pseudocephalozia (Hepaticae). J. Hattori Bot. Lab. 38: 665-701. Spruce, R. 1884-1885. Hepaticae amazonicae et andinae. Trans. Proc. Bol. Soc. Edinburgh 15: i-xi, 1-588. [Pp. 1-308 published in 1884, pp. 309-588 in 1885.] - - - . 1886. Voyage de Richard Spruce dans l'Amérique équatoriale pendant les années 1849-1864. Rev. Bryol. 13: 61-79. [Reprinted under the alternative title: Précis d'un voyage d'exploration botanique dans l'Amérique équatoriale, pour servir d'introduction provisoire a son ouvrage sur les hépatiques de 1'Amazone et des Andes, par Richard Spruce, p. 1-20.] Stotler, R. 1969. The genus Fruilania subgenus Fru/lania in Latín America. Nova Hedwigia 18: 397-555. Swails, L. F. 1970. The genus Parella in Latín America. Nova Hedwigia 19: 201-291. Thiers, B. M. 1993. A monograph of Pleurozia (Hepaticae: Pleuroziaceae). Bryologist 96: 517-554. Uribe M., J. & J. Aguirre C. 1995. Las especies Colombianas del género Symphyogyna (Hepaticae: Pallaviciniaceae). Caldasia 17: 429--458. Váiía, J. 1973. Studien über die Jungermannioideae (Hepaticae). 2. Jungermannia subg. Jungermannia. Folia Geobot. Phytotax. 8: 255-309. - - - . 1974. Studien überdie Jungermannioideae (Hepaticae). 4. Jungermannia subg. Plectoco/ea, subg. Solenostoma. Allgemeines. Süd- und mittelamerikanische Arten. Folia Geobot. Phytotax. 9: 179-208. - - - . 1976. Drei neue Gymnomitriaceae aus Südamerika. J. Hattori Bol. Lab. 41:411--417. - - - . 1980. Some new South and Central American hepalies. J. Hattori Bot. Lab. 48: 225-234. - - - . 1984. Anastrophyllum (Spruce) Steph. in Latín America-pre1iminary information. Proc. Third Meeting Bryol. Centr. East. Europe, Praha: 99-106. - - - . 1988. Cephalozia (Dum.) Dum. in Africa, with notes on the genus. Beih. Nova Hedwigia 90: 179-198. van der Grande, K. 1980. The genusJensenia Lindb. (Hepaticae). Proc. Kon. Ned. Akad. Wetensch. ser. C, 83: 271-278. van Reenen, G. B. A. 1982. High Andean species of Herbertus S. F. Gray. Lindbergia 8: 110-120. van Slageren, M. W. 1985. A taxonomic monograph ofthe genera Brachiolejeunea and Frullanoides. Meded. Bot. Mus. Herb. Rijksuniv. Utrecht 544: 1-309. Winkler, S. 1976. Die Hepaticae der Sierra Nevada de Santa Marta, Kolumbien. I: Terrestrische, epixyle und epipetrische Arten. Rev. Bryol. Lichénol. 42: 789-827. Yamada, K. 1988. The genus Radula from Cuba. J. Hattori Bot. Lab. 65: 379-390. Yuzawa, Y. 1991. A monograph ofsubgenus Chonanthelia of genus Frullania (Hepaticae) ofthe world. J. Hattori Bot. Lab. 70: 181-291.
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Vascular Plants James L. Luteyn
The checklist of vascular plants started with the lists ofC!eef(1981 b) and Vareschi (1970). Additions were made after scanning numerous monographs, revisions, and country or regional floras, such as Flora de Colombia, Flora of Ecuador, Flora Neotropica, and Flora de Venezuela. Then a very intensive and exhaustive family-by-family and genus-by-genus search for taxa (native ¡md naturalized) represented in the páramo flora was made at the United States National Herbarium (US) by myself and Mauricio Gavilanes, followed by lesser general searches by myself at AAU, NY, and U. Numerous specific herbaria (e.g., COL, CR, GB, GH, K, MER, MERF, MO, NY, S, VEN, etc.) were also consulted for particular families (or genera) when a specialist had worked there. Once the basic family or generic. lists were made, copies were sent to individual speciálists for comment, correction, and suggestion. Whenever possible, I have tried to include in this list only species for which I found specimen determinations made by recognized authorities. Norrnally I have not used species determined with the notations "aff." or "ve! aff.," and only rarely are species included when detennined with "cf." Furthermore, I have used a very conservative approach, including in the list only species for which I have seen at least one voucher from páramo (as herein defined), or when a literature citation or personal communication was considered by me to be extremely reliable. A bit of subjectivity was involved in the compilation of this checklist of páramo vascular plants. So me may say that the list does not include this or that species for which the !abe! clearly says "Páramo de XYZ." But it is sometimes very difficult to decide whether or not to in elude a species. In many cases, it is dangerous to include species for which the !abe! says only "páramo" and nothing else, often because collectors' concepts of páramo may differ. Severa! things must be remembered in this regard. First of all, páramo does not automatically include everything above the general3000 m elevation leve!. As has been stated in the Introduction of this book, montane forest may reach as high as 4300 m in the northern Andes. Second, the fact that a collectionlabel says "Páramo de XYZ" does not automatically mean that the plant was collected in a páramo habitat as herein defined. Among local people it is very common for the proper name "Páramo de XYZ" to include the full range of
low to very high elevations and montane forest to superpáramo-that is to say, the term "páramo" in itself is not enough of a criterion for including a plant in the checklist. Therefore, I have not included a species solely because the !abe! says "Páramo de XYZ" unless the !abe! also tells me something about the habitat (e.g., grass páramo), or I know the plant is a typical páramo plant, or I know the area from personal experience, or I know the collector and can be quite certain of what concept is used. On the other hand, so me may say that this or that genus or species found in this list ought not to have been included, because it occurs mostly in forest. This is where my own personal field experience (i.e., knowledge of the area) or definition of páramo is needed. Here, too, is where the difficulty of"What is subpáramo?" comes into play. For the purposes of this book and the checklist herein, subpáramo ineludes elements of small trees and shrubs in transition to grass páramo, and species ofthe isolated shrubtree islands or forest patches within open grass páramo. It may be debated whether or not the plants of the shrub-tree islands ought to be included in a páramo checklist, because they are usually forest species; but I have eh osen to include them just as I have included species of Polylepis forest. From the beginning ofthis project, my goal has been to presenta list of al! plants found in the páramo, whether from the more distinctive open grass páramo or superpáramo areas to the less distinctive, more transitional subpáramo zone that includes sorne forest elements. I ha ve not, however, included species of plants that are characteristic ofthe "southem Ecuadorean scrub" vegetation as defined by Harling (1979), which sorne call "subpáramo chaparral" and include in páramo vegetation, but which I consider a natural vegetation type distinct from páramo. The geographical and elevational ranges given in this checklist are only for species distributions within the páramo countries here considered (i.e., Costa Rica to northern Peru), not for total geographical and elevational ranges. This needs to be clarified with two hypothetical examples. Example 1: The geographical range given in the list for species A is CR, CO, EC, PE; however, it is al so found in PA and VE, but not in páramo there; and it is also found in Mexico and Bolivia, but these countries are outside the defined páramo range.
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Example 2: The elevational range given in the list for species A is 3100-3900 m; however, it also occurs in PA and VE, but at 2500-2700 m, which is not considered páramo; and it is found in Mexico and Bolivia at 1500-2100 m, but these elevations are not included because the countries are outside the defined páramo range. Because the data were gathered in this manner, 1 was not, unfortunate1y, able to calculate the total number of endemic páramo species. The taxa in this checklist are organized according to the rank of species, and 1 have not formal! y listed the ranks of subspecies or variety, when they occur. However, if a recognized species consists of one or more subspecies anci/or varieties, these are formally recognized at the end ofthe entry after the word "ineludes." Furthermore, certain taxa included in this checklist as synonyms may be recognized at another rank (perhaps as accepted species) in other publications, or, conversely, taxa recognized as species in other publications may be included as synonyms in this checklist. This is inevitable because species concepts often vary among specialists. Rather than trying to give complete synonymy, I have given only those synonyms in common or regional use or ofhistorical importance. Two genera that are cited as north-temperate elements reaching their southern limit at the Chirripó páramo in Costa Rica (Cleef & Rangel Ch., 1984; Cleef & Chaverri P., 1992), Helianthemum (Cistaceae) and Smilacina (Liliaceae), are not included in this checklist because I have not been able to locate herbarium specimens that clearly indicate a páramo habitat. As for the many species that are weeds (ruder¡1ls) or that have been cultivated in or near páramo and have subsequently escaped into páramo habitats, or those that are now naturalized-1 have included them in this list only when I have seen a herbarium voucher specimen. Many other species that have not yet become naturalized, or have escaped and are only rarely found in páramo, or are reported in the literature or by personal communication to me as occurring in páramo have not been included because 1 was unable to find voucher specimens in herbaria. Por discussions ofplants introduced to páramo regions, see Pérez, in press; Sauer, 1988; and Vareschi, 1970. The following species may be expected to occur in páramo following their escape from cultivation, but are not included in the list. Apiaceae Ciclospermum leptophy/lum (Pers.) F.Muell. Brassicaceae Arabis turrita L. and other spp. (native to Europe)
75
Brassica campes tris L. Raphanus raphanistrum L. (native to Europe, introduced and cultivated)
Caryophyllaceae Si/ene injlata Sm. (native to Europe)
Fabaceae Medicago lupulina L. (native to Europe, now cultivated, naturalized and used as forage) andM. denticulata Willd. Melilotus officinalis Willd. (native to Europe and Asia) U! ex europaeus L. (used as ornamental) Vicia faba L. (introduced as crop) and V. sativa L. (as forage)
Lamiaceae Mentha (cf. M. viridis L.)
Papaveraceae Papaver glaucum Boiss. & Hausskn. ex Boiss. (and perhaps P. somniferum L.) (native to Middle East)
Phytolaccaceae Phytolacca octandra L. (native to Asia)
Pinaceae Pinus radiata D.Don
Poaceae Avena sativa L. Hierochloe mexicana Benth. (native to Mexico ?) Hordeum muticum J.S.Presl (cultivated as forage) Lolium multiflorum Lam. and L. perenne L. (cultivated as forage) Melinis minutiflora P.Beauv. (native to Africa, introduced vi a Brazil, cultivated as forage) Pennisetum clandestinum Hochst. ex Chiov. (introduced from Africa, now escaped) Rhynchelytrum repens (Willd.) Hubb. [syn.: Tricholaena repens (Willd.) Hitchc.] (native to South Africa) Setaria parviflora (Poir.) Kerguélen [syn.: S. geniculata (Lam.) P.Beauv.] (cultivated as forage) Triticum aestivum L. (cultivated as crop)
Polygonaceae Rumex crispus L. (native to Europe)
The families given in the checklist are divided alphabetically within the larger artificial groups: fems and fern allies, gymnosperms, and angiosperms (including both monocots and dicots). The taxonomic system used is basically that of Cronquist (1981 ), with modifícations. Certain families such as Asteraceae and Poaceae, because they are so large, will always need additional work and updating. As mentioned abo ve, I feel most unsure about the Orchidaceae, because, in my opinion, they still need much basic taxonomic study, the nomenclature is confusing, they are very much undercollected at páramo elevations, and the
!
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herbarium specimens have labels with poor or little A. palyphyllum Bertol.; CR, VE; 2600-3400 m habitat data, are few in number, and were generally A. serra Langsd. & Fisch.; CO, PE; 100-3650 m unavailable for my study. I am indebted to Cal Dodson A. sessilifalium Desv.; CO, EC, PE; 2500-4000 m for sharing with me unpublished information from his A. squamasum L.; EC, PE?; 2100-3900 m A. triphyllum C.Presl; CO, EC, PE; 2800-4800 m database ofEcuadorean orchids. The list ends with literature citations of a strictly Cystapteris Bernh. taxonomic nature that have been cited in the check- C.fi'agilis (L.) Bernh. [syn.: C. diaphana (Bory) Blasdell]; list. These references are not meant.to be exhaustive; CR, CO, VE, EC, PE; 300-4500 m they are restricted to those with an emphasis on páramo taxa, those in which many páramo taxa are treated, and AZOLLACEAE those ofhistorical ami/or nomenclatura] importance. Azalla Lam. Finally, when questions arose concerning proper author citation or spellings of sorne species names, I A. caraliniana Willd.; VE?, EC; 3600-4100 m generally followed the TROPICOS database of the .. A.filiculaides Lam.; CO, VE, EC, PE; 3000-3600 m Missouri Botanical Garden, and for this help I am very BLECHNACEAE · much indebted to Peter M. J0rgensen. As mentioned abo ve, I ha ve tried to be consistent Blechnum L. in the way I recognize taxa in this checklist and ha ve B. auratum (Fée) R.M.Tryon & Stolze (syn.: B. buchtienii usually followed the taxonomy of one author or one Rosenst., B. calumbiense Hieron., B. ca/umbiense var. institutional system (e.g., US for Asteraceae); but new bagatense Hieron.); CR, PA, CO, VE, EC, PE; 19004000 m; includes subsp. auratum and subsp. ca/umbiense revisions, monographs, and floras appear every day. (Hieron.) R.M.Tryon & Stolze
B. binervatum (Poir.) C.V.Morton & Lellinger subsp.fragile R.M.Tryon & Stolze [syn.: B. fragile (Liebm.) C.V.Mortori & Lellinger]; CO; 1550-3500 m
CHECKLIST OF VASCULAR PLANTS Ferns and Fern Allies Ref.: Moran & Riba, 1995; Smith, 1985; Tryon & Stolze, 1989a, 1989b, 1991, .f992, 1993, 1994; Tryon & Tryon, 1982.
ADIANTACEAE Adiantum L. A. pairetii Wikstr.; EC, PE; 1200-4200 m A. raddianum C.Pres1; PE; 400-4000 m
ASPLENIACEAE Asplenium L. A. cuspidatum Lam.; CO, EC, PE; 1500-3950 m; includes var. cuspidatum and var.faeniculaceum (Kunth) Morton & Lellinger
A. haenkednum (C.Pres1) Hieron. (syn.: A. clada/eptan Fée); VE; 2900-4100 m
A. mananthes L. (syn.: A .. castaneum Schltdl. & Cham.); CO, VE, EC, PE; 1300-4900 m; intludes var. castaneum (Schltdl. & Cham.) Stolze and var. manan/hes A. aellgaardii Stolze; EC; 3400 m A. peruvianum Desv. (syn.: A.fi'agile C.Presl); CO, VE, EC, PE; 2750-4500 m
B. B. B. B.
cardatum (Desv.) Hieron.; CO, VE, PE; 700-4300 m lech/eri Mett.; CO, EC; 3000 m lima Rosenst.; CO, EC; 3200-3950 m /axense (Kunth) Hook. ex Salomon;_ CR, PA, CO, VE, · EC, PE; 1750-4100 m; includes var. /axense and var. stenaphyllum (Klotzsch) C. V.Morton B. accidenta/e L.; VE; 3400 m B. schamburgkii (Klotzsch) C.Chr.; CO, VE, EC, PE?; 1450-3700m
B. sprucei C.Chr.; EC; 3700 m
CYATHEACEAE Cyathea Sm. (syn.: Trichipteris C.Presl) [Note: All ofthe species of Cyathea listed here (except C. atahuallpa) are found in subpáramo woods or thickets or in paramillo and shrubby páramo (Tryon, 1976). !, myself, would not consider Cyathea a páramo genus, but in elude it here only on the basis ofTryon's paper.] Ref.: Tryon, 1976. C. atahuallpa (R.M.Tryon) Lellinger (syn.: Sphaerapteris atahua/lpa R.M.Tryon); PE; 2850-3450 m C. caracasana (Klotzsch) Domin var. baliviensis (Rosenst.) R.M.Tryon; CO; 1000-4200 m C. frigida (H.Karst.) Domin [syn.: Alsaphila frigida H.Karst.; Trichipteris frigida (H.Karst.) R.M. Tryon ]; CO, VE; 2800-3500 m C. fu/va (M.Mart~s & Galeotti) Fée; CO; 1300-4200 m C. pallescens (Soéliro) Domin; PE; 1250-3500 m C. suprastrigasa (H.Christ) Maxon; CR; 2000-3000 m
! .
1999]
PARAMOS
Trichipteris C.Pres1 (see Cyathea)
DENNSTAEDTIACEAE Histia;·teris (J.Agardh) J.Sm. H. incisa (Thunb.) J.Sm.; CO, VE, EC, PE; 1000-3750 m Hypolepis. Bemh. H. bogotensis H.Karst. (syn.: H. flexuosa Sodiro); CR, CO, EC; 2400-4050 m H. obtusata (C.Pres1) Kuhn ex Rieron.; CO, EC, PE; 28004600 m H. purdieana Rook.; CO, EC; 3900 m Pteridium G1ed. ex Scop. P. aquilinum (L.) Kuhn var. arachnoideum (Kau1f.) Brade [syn.: P. arachnoideum (Kau1f.) Maxon]; CO, PE; 4003000 m ·
DICKSONIACEAE Dicksonia L'Rér. D .. sellowiana Roo k. [syn.: D. karsteniana (K1otzsch) T.Moore, D. stuebellii Rieron.]; CO, PE; 2200-3450 m
DRYOPTERIDACEAE Athyrium Roth A. filix-femina (L.) Roth (syn.: A. dombeyi Desv., A. paramico/a L.D.Gómez); CR, CO, VE, PE; 2500-3550 m Dryopteris Adans. D. wallichiana (Spreng.) Ryl. [syn.: D. paleacea (Sw.) Rand.-Mazz., D. parallelogramma (Kunze) A1ston]; CO, VE, PE; 3200-4400 m Elaphoglossum Schott ex J.Sm. Ref.: Micke1, 1991, 1995. E. affine (M.Martens & Ga1eotti) T.Moore [syn.: E. revolvens (Kunze ex Kuhn) C.Chr.]; CR, CO, VE, PE; 3000-4050 m E. albescens (Sodiro) H.Christ; CO, VE?; 2500-3400 m E. anceps Micke1; VE; 3000 m E. andico/a (Fée) T.Moore; CO, VE?; 1200-3400 m E. andrea!1um R.Christ; CO; 3300 m E. atrorubens Mickel; VE; 3200-3600 m E. atrosquamatum Mickel; CO, VE, PE; 3500-4250 m E. bonapartei Rosenst.; EC; 4000? m E. camptplepis Mickel; PE; 3500 m E. cardiophyllum (Rook.) T.Moore; CO, EC; 3850-4000 m E. castaneum (Baker) Diels; CO, EC, PE; 1950-3450 m E. chrysopogon Mickel; VE; 1500-3300 m E. conspersum H.Christ; CR; 2500-3150 m E. corderoanum (Sodiro) H.Christ; CO; 4050 m E. cuspfdatum (Willd.) T.Moore; CO; 1200-3800 m E. delicatu/um Mickel; VE; 2500-3000 m E. deltoideum (Sodiro) H.Christ; CO; 3530 m E. dendricola (Baker) H.Christ; EC; 3050-3600 m E. deorsum (R.Karst.) Vareschi; VE; 2400-4000 m
77
E. dombeyanum (Fé.e) T.Moore & Rou1ston; VE, EC; 24003800m E. drewianum Micke1; VE; 3000-3200 m E. engelii (R.Karst.) H.Christ; CR, CO, VE, EC, PE; 28004600 m E. erinaceum (Fée) T.Moore; PE; 700-3400 m E.fournierianum L.D.Gómez; CR, PA; 900-3400 m E.funkii (Fée) T.Moore; CO; 3000-3400 m E.fiafuraceum (Mett. ex Kuhn) H.Christ; CR, PA; 900--3820 m E. glabellum J.E.Sm.; CO; 500-4100 m E. glossophyllum Rieron.; CO, EC; 2200-3750 m E. huacssaro (Ruiz) H.Christ; CR, CO, EC; 2900-3375 m E. inaequa/ifolium (Jenman) C.Chr. [syn.: Acrostichum sellowianum (K1otzsch ex Kuhn) T.Moore]; VE; 38003900 m E. latifolium (Sw.) J.Sm. [syn.: E. longifolium (C.Pres1) J.Sm.]; CO; 550-3800 m E. laxisquama Micke1; PE; 3350-3600 m E. leporinum L.D.Gómez; CR, PA; 3350-3550 m E. lindenii (Bory ex Fée) T.Moore; CR, CO, VE, EC, PE?; 3000-4500 m E. lingua (C.Pres1) Brack.; CO; 3000-4200 m E. /loense (Rook.) T.Moore; EC; 3200-3350 m E. luteynii Micke1; VE; 3000-3350 m E. mathewsii (Fée) T.Moore [syn.: E. hartwegii (Fée) T.Moore]; CO, VE, EC, PE; 3200-5100 m E. melancholicum Vareschi; CO, VE, PE; 2480-4050 m E. minutum (Poh1 ex Fée) T.Moore [syn.: E. gayanum (Fée) T.Moore,E.leptophyllum (Fée) T.Moore]; CR, PA, CO, VE, EC, PE; 2000-3900 m E. muscosum (Sw.) T.Moore; VE, EC; 2900-4100 m E. nigrocostatum Mickel; CO, EC; 1300-3750 m E. nivosum (Kunze) Micke1; VE, PE; 2600-3900 m E. odonto/epis Mickel; PE; 3200-4000'm E. ovatum (Rook. & Grev.) T.Moore; CO, VE, EC; 20004700 m E. paleaceum (Rook. & Grev.) S1edge; CO, EC, PE; 9503950m E. papillosum (Baker) H.Christ; VE, PE; 2000-3200 m E. petiolatum (Sw.) Urb.; CO; 1900-3380 m E. petiolosum (Desv.) T.Moore; CO, PE; 3050-3950 m E. pi/osius Micke1; CR, P A, CO, VE, PE; 2900-4600 m E. plicatum (Ca.v.) C.Chr.; CO, EC, PE; 3300-3450 m E. rimbachii (Sodiro) R.Christ (syn.: E. pichinchae H.Christ); EC, PE; 2300-4700 m E. rosenstockii H.Christ ex Rosenst.; EC, PE; 3400-3950 m E. ruficomus Micke1; PE; 3050-4200 m E. rupestre (R.Karst.) H.Christ; CO; 3800 m E. sporado/epis (Kunze ex Kuhn) T.Moore; VE; 750--3300 m E. squamipes (Rook.) T.Moore; VE, PE; 2400-3450 m E. stramineum (Mett.) T.Moore; CO; 3400-3550 m E. tabanense André ex H.Christ; CO; 3050-3725 m E. tachirense Micke1; VE; 3100-3300 m E. tectum (Rumb. & Bonpl. ex Willd.) T.Moore; EC; 19003650 m E. tenuifolium (Liebm.) T.Moore; CO; 3950 m E. vagans (Mett.) Rieron.; CO; 3650-3700 m E. vulcanicum H.Christ; EC; 2500-3500 m E. yatesii (Sodiro) R.Christ; CO, EC; 3600-4500 m
.-L 1
. !
78
MEMOIRS OF TRE NEW YORK BOT ANICAL GARDEN
Po/ystichum Roth P. aculeatum (L.) Schott [syn.: P. coch/eatum (Klotzsch) Rieron.]; CO, VÉ, EC; 3100--4300 m P. montevidense (Spreng.) Rosenst. var. nudicau/e (Rosenst.) R.M.Tryon (syn.: P. nudicaule Rosenst.); CO, VE, PE; 3200--4400 m P. muricatum (L.) Fée; EC; 1500-3650 m P. orbiculatufn (Desv.) J.Remy & Fée var. orbiculatum [syn.: P. polyphyllum (C.Pres1) C.Pres1]; CR, CO, VE, EC, PE; 3000--4900 m ) P. pumilisMaxon; CO, EC; 3500-3700 m P. pycno/epis (Kunze ex K1ot¡¡sch) T.Moore [syn.: P. gelidum (Kunze ex K1otzsch) Fée]; CO, VE, EC, PE; 3000--4400 m P. sodiroi H.Christ; CO, EC; 3220-3600 m P. speciosissimum (A.Braun ex Kunze) Cope!. [syn.: Plecaso rus speciosissimum (A.Braun ex Kunze) T.Moore]; CR; 2500-3500 m P. ta/amancanum Barrington; CR, PA; 3100-3500 m Woodsia R.Br. W. montevidensis (Spreng.) Rieron. [syn.: W. crenata (Kunze) Rieron.]; CO, PE?; 3200--4400 m
EQUISETACEAE Equisetum L. E. bogotense Kunth; CO, VE, EC, PE;. 3000-3800 m
GLEICHENIACEAE Gleichenia Sm. (syn.: Sticherus C.Pres1) G. revo/uta Kunth [syn.: S. revolutus (Kunth) Ching]; EC, PE; 1750-3500 m G. rubiginosa Mett. [syn.: S rubiginosus (Mett.) Nakai]; PE; 11 00--4200? m G. simplex (Desv.) Rook. [syn.: S. simplex (Desv.) Ching]; CO, EC, PE; 2700--4000 m Sticherus C.Pres1 (see Gleichenia)
[VOL. 84
po/yanthos (Sw.) Sw.; PA, CO, VE; 3100--4450 m pulchel/um Schltd. & Cham.; CR; 3350-3550 m ruizianum (K1otzsch) Kunze; CO; 2800 m si/iquosum H.Christ; CR; 3100-3'400 m tegularis (Desv.) Proctor & Lourteig (syn.: H. e/egantulum Bosch); PA, CO, PE; 3100--4000 m H. trichophyllum Kunth; CO, VE, EC, PE; 2200--4450 m H. tunbrigense (L.) Sm.; CR, VE, EC; 2600--4000 m · H. undu/atum (Sw.) Sw. var. undulatum; CR, CO, EC?; 1500--4100m H. H. H. H. H.
ISOETACEAE Isoi!te1- L. (syn.: Stylites Amstutz) Rd.: Fuchs-Eckert, 1982; Rickey, 1985. l. a~dico/a (Amstutz) L.D.Gómez (syn.: Stylites andicola Amstutz); CO; 3600 m 1. andina Spruce ex Rook. (syn.: l. triquetia A.Braun); CO, VE, EC; 3400--4300 m l.bisch/erae R.P.Fuchs; CO; 32Ó0-3600 m l. boliviensis Weber; PE; 4100-5000 m l. boyacensis R.P.Fuchs; CO; 3400--4000 m l. c/eejli R.P.Fuchs; CO; 3750--4250 m l. colombiana (T.C.Pajmer) H.P.Fuchs (syn.: l. /ech/eri Mett. var. colombiana T.C.Pa1mer); CO; 3900 m 1. dispara Rickey; PE; 3300 m l. ecuadoriensis Asp!.; EC; 4240 m l. hewitsonii Rickey; PE; 3700 m 1. karstenii A.Braun; CO, VE; 3400--4100 m 1. killipii C.V.Morton; CO, VE, EC; 3200--4250 m l. lech/eri Mett. (syn.: l. socia A.Braun, /. g/acia/is Aspl.); CO, VE, PE; 3300--4800 m 1. novo-granadensis R.P.Fuchs (syn.: 1. dichotoma L.E.Mora & W.Ragemann); CO, EC; 3250--4300 m l. pa/meri R.P.Fuchs; CO, VE, EC; 3350--4450 m 1. ramboi Rerter; CO; 3800 m l. rimbachiana H.P.Fuchs; EC; 4200 m l. storkii T.C.Pa1mer (syn.: l. tryoniana L.D.Gómez); CR, PA; 3100-3800 m Stylites Amstutz (see lsoi!tes)
HYMENOPHYLLACEAE Hymenophyllum Sm. (syn.: Sphaerocionium C.Pres1) H. amabile C.V.Morton; VE, EC, PE; 2600--4200 m H. cristatum Rook. & Grev.; EC; 3500-3800 m H.ferax Bosch; CO; 2400-3300 m H.fucoides (Sw.) Sw. (syn.: H. ca/odictyon Bosch); CO, VE, PE; 1700-3700 m H. gol/meri Bosch; CO, EC; 3800-3900 m H. jamesonii Roo k.; CO, EC; 3100-3900 m H. karstenianum J.W.Sturm; PE; 3100 m H. molle C.V.Morton; CO; 3500 m H. multialatum C.V.Morton; PE; 200--4200? m H. myriocarpum Rook. var. endiviifolium (Desv.) Sto1ze (syn.: H. andinum Bosch, H. nigrescens Liebm., H. multijlorum Rosenst.); PA, CO, VE, EC, PE; 2000--4200 m H. peltatum (Poir.) Desv.; EC; 2800--4700 m H. plumier! Roo k. & Grev.; CO, PE; 3000-3500 m
LOPHOSORIACEAE Lophosoria C.Pres1 L. quadripinnata (J.F.Gme!.) C.Chr. var. contracta (Rieron.) R.M.Tryon & A.F.Tryon; CO, EC, PE; 2800-3600 m
L YCOPODIACEAE Ref.: 0llgaard, 1987, 1988, 1989, 1992, 1995. Huperzia Bernh. (syn.: Urostachys Rerter) H. andina (Rosenst.) Ro1ub; PE; 3300-5000 m H. arcturii (Rerter) C.V.Morton; CO, VE; 3200-3500 m H. ascendens (Rerter ex Nesse1) Rolub; EC; 3370--4050 m H. attenuata (Spring) Trevis.; CR, EC, PE; 3100--4000 m H. austroecuadorica B.0llg.; EC; 2500-3400 m H. binervia (Rerter) B.0llg.; PE; 2000-331 O m
1999]
PARAMOS
H. brevifolia (Grev. & Hook.) Ho1ub (syn.: H. goudotii Herter); CR, CO, EC, PE; 2700-4200 m H. cape/lae (Herter) Holub [syn.: Lycopodium capellae (Herter) C.V.Morton]; CO, VE, EC, PE; 3050-4300 m H. catacachiensis (Herter ex Nessel) B.011g.; CO; ? m H. chiricana (Maxon) Holub; PA; 3400 m H. colanensis B.01lg.; PE; 3140-3600 m H. columnaris B.01lg.; EC, PE?; 3800-4000 m H. compacta (Hook.) Trevis.; EC; 3000-3550 m H. crassa (Humb. & Bonpl. ex Willd.) Rothm. (syn.: Lycopodium erythraeum Spring); CR, PA, CO, VE, EC, PE; 3500-4700 m; in eludes vars. crassa, gelida B.01lg., and manus-diaboli B.0llg. H. cruenta (Spring) Rothm.; GO, VE; 3100-3700 m H. cumingii (Nessel) Holub; EC; 3350-4300 m H. dianae (Herter) B.01lg.; CO; 3650 m H. espinosana B.011g.; EC; 3000-3400 m H. eversá (Poir.) B.01lg. (syn.: Lycopodium ecuadoricum Herter); CR, PA, CO, PE; 3000-3700 m H. hartwegiana (Spring) Trevis.; CO, VE, EC; 2700-3850 m H. hasta/a B.01lg.; EC; 3000-3450 m H. bippuridea (H.Christ) Holub; CO, VE, EC, PE; 2400. 3600 m H. hoffmannii (Maxon) Rolleri & Deferrari; CR, PA; 20003600 m H. hohenackeri (Herter) Holub; CO, EC; 3400-4400 m H. hypogaea B.011g.; CO, EC, PE; 3000-4450 m H. hystrix (Herter) Holub; CO, EC; 3100-4100 m H. cf. innocentium (Herter) Holub; CO?, VE; 3300-4250 m H. kuesteri\(Nesse!) B.011g.; EC, PE; 2750-3600 m H. lignosa (Herter) Holub [syn.: H. serpentiformis (Herter) Rolleri & Deferrari]; CO; 3100-3400 m . H. lindenii (Spring) Trevis.; CO, EC; 3500-4200 m H. llanganatensis B.01lg.; EC; 3600-4000 m H. loxensis B.011g.; EC; 2500-3200 m H. macbridei (Herter) B.011g.; EC, PE; 2550-3650 m H. m:Yf'tuosa (Spring) Trevis.; CO, VE; 3000-3500 m H. ocanana (Herter) Holub (syn.: Lycopodium moritzii Grev.); CO, VE; 3000-4000 m H. pflanzii (Nessel) Rolleri & Defarrari; CR; 3100-3200 m H. phylicifolia (Desv. ex Poir.) Ho1ub; CO, EC, PE; 23003700 m H. polydactyla B.01lg.; EC; 3800-4200 m H. polylepidetorum B.01lg.; EC; 3650-4000 m H. rejlexa (Lam.) Trevis.; CR, CO, VE, EC, PE; 900-3400 m H. riobambensis (Herter) B.01lg. [syn.: H. castoris (Herter) C.V.Morton]; CO, VE; 3100-3400 m H. rosenstockiana (Herter) Holub [syn.: H. sacra/ea (Herter) Rolleri & Deferrari]; CO, EC, PE; 3000-3700 m H. rufescens (Hook.) Trevis.; CO, VE, EC; 3200-4200 m H. sagasteguiana B.01lg.; PE; 3900-3950 m H. saururus (Lam.) Trevis.; PE; 3300-4500 m H. scabrida B.01lg.; EC; 3800-4000 m H. schlimii (Herter) B.011g.; CO; 3250-3300 m H. schmidtchenii (Hieron.) Holub; CO; 3300-3750 m H. sellifolia B.01lg.; CO, EC, PE; 3100-4200 m H. subulata (Desv. ex Poir.) Holub; CR, PA, CO, EC, PE; 2850-3450 m
H. H. H. H. H. H. H. H. H. H.
79
talamancana B.01lg.; CR, PA; 3000-3400 m talpiphila B.01lg.; EC; 3400-4150 m taxifolia (Sw.) Trevis.; PA, CO; 3100-4330 m tetragona (Hook. & Grev.) Trevis.; CO, EC, PE; 37004200m transilla (Sodiro ex Baker) Holub; CO, EC; 3700-4250 m ulixis (Herter) Holub; CO; 3700 m urbanii (Herter) Holub; EC; 2800-3800 m venezuelanica (Herter) Holub (syn.: Lycopodium schneei Vareschi); CO, VE; 3600-4!'00 m weberbaueri (Nessel) Ho1ub; EC, PE; 3000-3600 m weddellii (Hieron. & Herter ex Nessel) Holub; EC, PE; 2600-3900 m
Lycopodiella Holub L. alopecuroides (L.) Cranfill; PE; 2400-3450 m L. caroliniana (L.) Pic.Serrn. var. meridionalis (Underw. & Lloyd) B.01lg. & Windisch; PE; 1350-3450 m L. matthewsii (Hook.) Holub; CO, VE, EC, PE; 2950-3650 m L. pendulina (Hook.) B.011g.; CR, CO, VE, EC, PE; 22003600m Lycopodium L. L. clavatum L. subsp. contiguum (Klotzsch) B.01lg. (syn.: L. contiguum Klotzsch); CR, PA, CO, VE, EC, PE; 29004100 m; includes var. aristatum (Humb. & Bonpl. ex Willd.) Spring L.jussiaei Desv. ex Poir.; CR, CO, VE, EC, PE; 2900-3900 m L. magellanicum (P.Beauv.) Sw. [syn.: L. spurium Wílld.]; CR, CO, VE, EC, PE; 2750-4200 m L. thyoides Humb. & Bonpl. ex Willd. (syn.: L. complanatum L. var. tropicum Spring); CR, CO, VE, EC, PE; 20004000m L. vestitum Desv. ex Poir.; EC, PE; 2400-3800 m
MARSILEACEAE Pilularia L. P. americana A.Braun; VE; 3300-3500 m P. mandonii A.Braun; CO, VE; 4000-4100 m
OPHIOGLOSSACEAE Bollychium Sw. B. schaffneri Underw. [syn.: B. ternatum (Thunb.) Sw.]; CO, VE, PE; 2350-3850 m B. virginianum (L.) Sw.; PE; 1800-3450 m Ophioglossum L. O. crotalophoroides Walt.; CO, VE, EC, PE; 600-4400 m O. reticulatum L. (syn.: O. petiolatum Hook.); VE; 400-3800 m
PLAGIOGYRACEAE Plagiogyria (Kunze) Mett. Ref.: Lellinger, 1971. P. semicordata (C.Presl) H.Christ (syn.: P. costaricensis Mett. ex Kuhn); CR, CO, VE, EC, PE; 1800-3600 m
MEMOIRS OF THE NEW YORK BOTANICAL GARI)EN
80
POLYPODIACEAE Campyloneurum C.Presl C. amphostenon (Kunze ex Klotzsch) Fée (syn.: C. irregulare Lellinger; Po/ypodium amphostenon Kunze ex K1otsch, P. angustifo/ium Sw. var. amphostenon (Kunze ex Klotzsch) Baker, P. /eucorhizon K1otzsch); CR, PA, CO, VE, EC, PE; 2650--4200 m C. angustifolium (Sw.) Fée [syn.: C. ensifo/ium (Willd.) J.Sm.; Po/ypodium angustifolium Sw.]; CO, VE, EC, PE; 2000--4500 m C. cochense (Hieran.) Ching; CO; 3-300-3790 m C. densifolium (Hieran.) Lellinger; CO, VE, PE; 3000-3600 m C.fascia/e (Humb. & Bonpl. ex Willd.) C.Presl; VE; 30003700m C. remotifolium (I;Iieron.) Lellinger; CO, VE, EC; 34003800 m C. solutum (K1otzsch) Fée; CO, EC, PE; 3750--4000 m Ceradenia L.E.Bishop Ref.: Bishop, 1988. C. aulaeifolia L.E.Bishop ex A.R.Sm.; CR, PA; 3100-3350 m C. farinosa (Hook.) L.E.Bishop [syn.: Grammitis farinosa (Hook.) C.V.Morton]; CO, EC; 3650--4050 m C. intricata (C.V.Morton) L.E.Bishop ex A.R.Sm. [syn.: Grammitis intricata C.V.Morton]; CO, EC; 3800-3900 m
C. mayoris (Rosenst.) L.E.Bishop (syn.: Polypodium mayoris Rosenst.); CO; 3400-3700 m C. ph/oiocharis L.E.Bishop; PA; 3100-3300 m C. semiadnata (Hook.) L.E.Bishop [syn.: Grammitis semiadnata (Hook.) C.V.Morton]; CO, EC; 3050--4000 m Cochlidium Kau1f. Ref.: Bishop, 1978. C. serru/atum (Sw.) L.E.Bishop (syn.: Polypodium dua/e Maxon); CO; 3360-3470 m Ctenopteris Kunze (see Grammitis) Enterosora Baker Ref.: Bishop & Smith, 1992. E. asplenioides L.E.Bishop (syn.: Gymnogramma sinuata T.Moore ex Baker); CO; 3000 m Grammitis Sw. (syn.: Ctenopteris Kunze) (see a1so synonyms under Ceradenia, Lellingeria, Melpomene, Micropo/ypodium, Terpsichore, and Zygoph/ebia) Ref.: Morton, 1967. G. leptopoda (C.H.Wright) Copel.; CO; 2000-3400 m G. paramicolq,L.E.Bishop; CO, PE; 2750-3500 m
[VOL.
~4
myosuroides Sw.; Xiphopteris jamesonii Hook.]; VE, EC; 3100-3450 m L. pseudocapillaris (Rosenst.) A.R.Sm. & R. C. Moran [syn.: Grammitis pseudocapillaris (Rosenst.) C.V.Morton]; EC; 2200-3550 m L. tmesipteris (Cope!.) A.R.Sm. & R. C. Moran [syn.: Grammitis tmesipteris (Cope!.) F.Seym.]; CR; 2700-3300 m Me/pomene A.R.Sm. Ref.: Smith & Moran, 1992. M. andico/a (Sto1ze) A.R.Sm .. & R.C.Moran (syn.: Grammitis andico/a Sto1ze); CO, PE;? m M. anfractuosa (Kunze ex K1otzsch) A.R.Sm. & R.C.Moran [syn.: Grammitis anfi"actuosa (Kunze ex K1otzsch) Proctor]; CO; 2200--4300 m M.'assurgens (Maxon) A.R.Sm.·& R.C.Moran [syn.: Grammitis assurgens (Maxon) C.V.Morton)]; CO, EC; 2900-3700 m M e f. firma (J.Sm.) A.R.Sm. & R.C.Moran [syn.: Grammitis firma (J.Sm.) C.V.Morton]; VE, PE; 1200-3800 m Mjlabelliformis (Poir.) A.R.Sm. & R.C.Moran [syn.: Polypodium rigescens Bory ex Willd.; Grammitis jlabel/i./órmis (Po ir.) C.V.Morton, G. rigescens (Bory ex Willd.) Lellinger]; PA, CO, VE, EC, PE; 3000--4800 m M moni/iformis (Lag. ex Sw.) A.R.Sm. & R.C.Moran [syn.: Grammitis moniliformis (Lag. ex Sw.) Proctor]; CR, PA, CO, VE, EC, PE; 3100--4600 m M. peruviana (Desv.) A.R.Sm. & R.C.Moran [syn.: Grammitis peruviana (Desv.) C.V.Morton]; CO, VE, EC; 3000--4450 m M. pi/osissima (M.Martens & Ga1eotti) A.R.Sm. & R.C.Moran [syn.: Grammitis pilosissima (M.Martens & Ga1eotti) C.V.Morton; Polypodium acrodontium Fée]; CR, CO, VE, PE?; 2900--4200 m M. pseudonutans (H.Christ & Rosenst.) A.R.Sm. & R. C. Moran [syn.: Grammitis pseudonutans (H.Christ & Rosenst.) C.V.Morton]; CO, EC, PE; 3000--4000 m M sodiroi (H.Christ & Rosenst.) A.R. Sm. & R. C. Moran· [syn.: Polypodium sodiroi H.Christ & Rosenst.; Grammitis sodiroi (H.Christ & Rosenst.) C.V.Morton; Xiphop- · teris sodiroi (H.Christ & Rosenst.) Crabbe]; EC; 4200 m M. xiphopteroides (Liebm.) A.R.Sm. & R.C.Moran [syn.: Polypodium rigens Maxon, P. xiphopteroides Liebm.; Grammitis rigens (Maxon) Proctor, G. xiphopteroides (Liebm.) A.R.Sm.]; CO, VE, EC, PE?; 3000-3800 m Micropolypodium Hayata Ref.: Smith, 1992. M aphe/olepis (C.V.Morton) A.R.Sm. (syn.: Grammitis aphelolepis C.V.Morton); CO, EC; 2400-3500 m Niphidium J.Sm.
Lellingeria A.R.Sm. & R. C. Moran Ref.: Smith et al., 1991. L. humilis (Mett.) A.R.Sm. & R.C.Moran [syn.: Grammitis humilis (Mett.) Lellinger]; CO; 2700-3400 m L. myosuroides (Sw.) A.R.Sm. & R.C.Moran [syn.: Grammitisjamesonii (Hook.) C.V.Morton; Po/ypodium
.N. crassifolium (L.) Lellinger(syn.: Po/ypodium crassijloium
L.); EC, PE; 1000-3400 m Pec/uma M.G.Price P. absidatum (A.Evans) M.G.Price (syn.: Po/ypodium absidatum A.Evans); CO; 3800--4100 m
1999]
PARAMOS
P. curvans (Mett.) M.G.Price (syn.: P¡¿lypodium curvans Mett.); CO; 3400-3850 m P. eurybasis (C.Chr.) M.G.Price (syn.: Po/ypodium eW)'basis C.Chr.); CO, PE; 2500-3600 m /.
'
P/eopeltis Humb. & Bonpl. ex Willd. P. macrocarpa (Bory ex Willd.) Kaulf. (syn.: Polypodium /anceolatum L.); VE; 3000-3900 m ·' Polypodium L. P. chirripoense Lellinger; CR; 3100-3400 m P. chryso/epis Hook. [syn.: Microgramma chryso/epis (Hook.) Crabbe]; PE; 3200-3500 m P.fimbriatum Maxon; CO; 1500-3300 m P. gil/iesii C.Chr.; CO; 3000-3700 m P. /asiopus K1otzsch; CO, VE; 800-3700 m P. macrolepis Maxon; CR; 3100-3400 m P. mindense Sodira; CO, EC; 3850 m ,P. monosorum Desv.; CO, EC; 3000~200 m P. murorum Hook.; CO, VE, EC; 2800~100 m P. pycnocarpum C.Chr. var. buchtienii (H.Christ & Rosenst.) R.M.Tryon & Sto1ze (syn.: Pplypodium buchtienii H.Christ & Rosenst.); PE; 800-3650 m P. subandinum Sodira; CO, EC; 3000-3450 m .P. wiesbauerii Sodiro; CO; 3300-3600 m
\ Terpsichore A.R.Sm. Ref.: Smith, 1993. T. athyrioides (Hook.) A.R.Sm. [syn.: Grammitis athyrioides (Hook.) C.V.Morton]; PE; 3400-3550 m T. cultrata (Bory ex Willd.) A.R.Sm. [syn.: Grammitis cultrata (Bory ex Willd.) Practor]; EC, PE; 1800-3400 m T. heteromorpha (Hook. & Grev.) A.R.Sm. [syn.: Grammitis heteromorpha (Hook. & Grev.) C.V.Morton]; CO, VE, EC; 3400~600 m T.jamesonioides (Fée) A.R.Sm. [syn.: Grammitisjamesonioides (Fée) C.V.Morton]; CO, VE; 2900-3750 m T. lanigera (Desv.) A.R.Sm. [syn.: Grammitis /anigera (Desv.) C.V.Morton]; CO, EC, PE; 1000~100 m T. /eucosticta (J.Sm.) A.R.Sm. [syn.: Grammitis /eucostricta (J.Sm.) C.V.Morton]; CO; 3200-3400 m T. semihirsuta (Klotzsch) A.R.Sm. [syn.: Grammitis semihirsuta (Klotzsch) C.V.Morton]; CO, EC; 2200~000 m T. subscabrum (Klotzsch) A.R.Sm. [syn.: Grammitis pichinchensis (Hieran.) C.V.Morton, G. subscabrum (Klotzsch) C.V.Morton; T. pichinchensis (Hieran.) A.R.Sm.]; EC; 3100-3500 m T. variabilis (Mett. ex Kuhn) A.R.Sm. [syn.: Po/ypodium variabile Mett. ex Kuhn; Grammitis varia bilis (Mett. ex Kuhn) C.V.Morton]; CO, EC; 3700~350 m Xiphopteris Kaulf. {see Grqmmitis) Zygophlebia L.E.Bishop Ref.: Bishop, 1989. Z. eminens (C.V.Morton) L.E.Bishop (syn.: Grammitis eminens C.V.Morton); EC; 2750-3350 m
81
PTÉRIDACEAE Cheilanthes Sw. hirsuta Link; CO; 3000 m /endigera (Cav.) Sw.; VE, EC; 2600-3600 m margina/a Kunth; CR, CO, VE, EC; 1500~000 m myriophylla Desv.; PE; 1550-3250 m scariosa (Sw.) Presl.; PE; 3400~000 m
C. C. C. C. C.
Eriosorus Fée Ref.: Tryon, 1970. E. accrescens A.F.Tryon; PE; 2900-3500 m E. cheilanthoides (Sw.) A.F.Tryon; EC, PE; 2300~000 m E. e/ongatus (Hook. & Grev.) Cope!.; EC, PE; 230~ 100m E. ewanii A.F.Tryon; CO, EC; 3350-3950 m E. flabellatus (Hook. & Grev.) Cope!.; CO, PE; 3400 m [Note: Thought to be a hybrid between E. chei/anthoides andE. sp. (Tryon & Stolze, 1989b).] E.jlexuosus (Kunth) Cope!. var.jlexuosus [syn.: E. villosu/us (Maxon) Scamman]; CR, PA, CO, VE, EC; 220~200 m E. hirsutulus (Mett.) A.F.Tryon (syn.: Gymnogramma karstenii Mett.); CO; 2800~200 m E. hispidulus (Kunze) Vareschi var. hispidulus; VE; 30003100m E. lindigii (Mett.) Vareschi; CO; 2800-3350 m E. /ongipetiolatus (Hieran.) A.F.Tryon; CO, EC; 3300--3900 m E. novagranatensis A.F.Tryon; CO, EC; 3000~050 m E. rufescens (Fée) A.F.Tryon; VE, EC, PE; 2300-3700 m E. setu/osus (Hieran.) A.F.Tryon; CO, EC; 3000-3750 m E. warscewiczii (Mett.) Cope!.; CR, CO; 3200-3550 m Jamesonia Hook. & Grev. Ref.: Tryon, 1962. J. a/stonii A.F.Tryon; CR, PA, CO, EC, PE; 3300~200 m J. auricu/ata A.F.Tryon; VE; 2600-3200 m J. b/epharum A.F.Tryon; CO, EC; 1500-3400 m J. bogotensis H.Karst.; CO, VE; 2950~400 m J. canescens Kunze; CO, VE; 2600~400 m J. cera cea Maxon; CO; 3100-3200 m J. cinnamomea Kunze; CO, EC, PE; 3100-5000 m J. cuatrecasasii A.F.Tryon; CO, VE; 3500~400 m J. goudotii (Hieran.) C.Chr.; CO, VE, EC, PE; 300~800 m J. imbrica/a (Sw.) Hook. & Grev.; CO, VE, EC, PE; 30004400 m; includes var. glutinosa (H.Karst.) A.F.Tryon, var. imbrica/a & var. meridensis A.F.Tryon J. laxa (Mett. ex Kuhn) Die1s; VE; 2900~450 m J. peruviana A.F.Tryon; PE; 2800~300 m J. pulchra Hook. & Grev.; CO, VE?, EC, PE; 3000~250 m J. robusta H.Karst.; CO, EC; 2700-3600 m J. rotundifolia Fée; CR, CO, EC, PE; 3000~200 m J. sca/aris Kunze; VE?, PE; 2700~300 m J. scammanae A.F.Tryon; CR, PA, CO, EC; 2750~400 m J. verticalis Kunze; CO, EC; 3000-3600 m Nephopteris Lellinger Re f.: Lellinger, 1966. N. maxonii Lellinger; CO; 2900-3200 m
·..
82
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
. •Pel/aea Link P. ternifolia (Cav.) Link var. ternifolia; CQ, PE; 1800-4600 m Pityrogramma Link P. cluysoconia (Desv.) Maxon ex Domin [syn.: P. ornithopteris (Kiotzsch) Maxon ex R.Knuth]; CR, PA, CO, VE; 2500-3850 m P. tartarea (Cav.) Maxon; VE; 1200-3000.m Pteris L. P. longipetiolulata Lellinger; CO; 3200-3500 m P. muricata Hook.; CO; 1700--4000 m
SELAGII'!ELLACEAE Selaginella P.'Beauv. S. carinata R.M.Tryon; EC; 3500 m S. cavifolia A.Braun; CO, EC; 2100-3680 m
THEL YPTERIDACEAE Thelypteris Schmidel T. arenosa A.R.Sm.; CO; 3000 m T. atropurpurea Hieron.; CO; 3800--4200 m T. caucaensis (Hieron.) Alston; CR, CO, EC, PE; 33004300 m T. cheilanthoides (Kunze) Proctor; VE; 2500-3000 m T. euthythrix A.R.Sm.; EC; 3400-3450 m T. gomeziana A.R.Sm. & Lellinger; CR; 3000-3300 m T. minutula C.V.Morton; CO, EC; 3550--4500 m T. perstrigosa (Maxon) Ching; CO; 3000-3300 m T. rigescens (Sodiro) A.R.Sm.; EC; 2800-3900 m T. rosenstockii (C.Chr.) R.M.Tryon; EC; 3000-3300 m T. rudis (Kunze) Proctor; CO; 1300--4000 m T. subtilis A.R.Sm.; EC; ·3200-3350 m T. supina (Sodiro) A.R.Sm.; EC; 3100-3550 m
VITTARIACEAE Re f.: Crane, 1997. Radiovittaria (Benedict) E.H.Crane R. ruiziana (Fée) E.H.Crane (syn.: Vittaria ruiziana Fée); PE; 1800-3650 m
[VOL. 84
Angiosperms Ref.: Brako & Zarucchi, 1993; Davidse et al., 1994; J0rgensen & Ulloa U., 1994; Macbride et collab., 1936-present.
ACANTHACEAE Stenandrium Nees S. dulce (Cav.) Nees; CO, PE; 3100-3400 m
ALSTROEMERIACEAE Bomarea Mirb. B. acutifolia (Linh. & Otto) Herb,; CR, CO; 3200-3550 m B. ambigua Sodiro; EC; 3780 m B. anceps (Ruiz & Pav.) Herb.; PE; 3400-3500 m B. angustipetala (Benth.) Baker; CO; 3500 m B. cf. angustissima Killip; PE; 3500 m B. caldasii (Kunth) Asch. & Graebn.; CO, EC; 2100-4200 m B. chimboracensis Baker; EC; 3800 m B. crassifolia Baker; CO; 3200-3500 m B. dulcis (Hook.) Beauverd; PE; 3200--4500 m B. glaucescens (Kunth) Baker; EC, PE; 2900--4400 m B. hazenii Killip; CO; 3400 m B. hirsuta (Kunth) Herb.; CR, PA, CO, EC; 2000-3540 m B. linifo/ia (Kunth) Baker; CO, EC; 2500-3800 m B. multiflora Mirb.; VE; 4050--4100 m B. nervosa (Herb.) Baker; PE; 3100-3400 m B. pauciflora (Kunth) Herb. (syn.: B. holtonii Hochr.); CO, VE; 2500-3500 m B. purpurea (Ruiz & Pav.) Herb.; PE; 2650-3500 m B. setacea (Ruiz & Pav.) Herb.; CO, VE, EC, PE; 2400-4200 m B. stenopetala Baker; CO; 3000-3200 m B. cf. subspicata Sodiro; PE; 3200-3400 m B. tomentosa (Ruiz & Pav.) Herb.; CO; 2900-3450 m B. torta (Kunth) Herb.; PE; 2800-3600 m
AMARANTHACEAE Alternanthera Forssk. A. lupulino Kunth; PE; 3800-3850 m A. macbridei Stand!.; PE; 2500-3500 m
AMARYLLIDACEAE Vittaria Sm. V. graminifo/ia Kaulf.; VE, PE; 1200--3150 m
Gymnosperms Ref: Brako & Zarucchi, 1993.
Stenomesson Herb. S. aurantiacum (Kunth) Herb.; EC, PE; 3300-3400 m S. variegatum (Ruiz & Pav.) J.F.Macbr.; PE; 3000-3500 m
APIACEAE Ref.: Mathias & Constance, 1976b. Apium L. (see NiphogetÓn)
EPHEDRACEAE Ephedra L. E. americana Humb. & Bonpl. ex Willd. (syn.: E. andina Poepp.; EC, PE; 3800--4600 m E. rupestris Benth.; EC, PE; 3500--4500 m
Arracacia Bancr. A. atropurpurea (Lehm.) Benth. & Hook.f. ex Hemsl.; CR?, PA; 3500--4000 m A. elata H.Wolff (syn.: A. pennel/ii Constance); CO, VE?, EC, PE; 3400--4100 m
·\
1999]
PARAMOS
A. maschata (Kunth) DC.; CO, EC; 3300-3400 m; possibly cultivated A. talucénsis (Kunth) Hemsley var. multifida (S.Wats.) Jy!athias & Constance; CO; 3250-3850 m A. vaginata Cou1t. & Rose; VE; 2000--4200 m Azorella Lam. Ref.: Martínez, 1993. A. aretiaides (Spreng.) DC.; CO, EC; 3500--4500 m A. hilaba (Schltdl.) Wedd. (syn.: A. lehmannii Hieron.); CR, EC, PE; 3500--4400 m A. carymbasa (Ruiz & Pav.) Pers.; CO, EC, PE; 3600--4450 m A. crenata (Ruiz & Pav.) Pers.; CO, VE,EC, PE; 3300--4200 m . A'. cuatrecasasii Mathi¡¡s & Constance; CO, VE; 3000-3800 m A. ecuadorensis Domin; EC; 3700--4600 m A. julianii Mathias & 'Constance; VE; 3250--4550 m A. multifida (Ruiz & Pav.) Pers.; CO, EC, PE; 3500--4300 m A. pedunculata (Spreng.) Mathias & Constance; CO, EC; . 3550--4650 ni Bowlesia Ruiz & Pav. B. !abata Ruiz & Pav.; EC, PE; 3450--4100 m B. sodiroana H.Wo!:l'f; EC; 3650 m Cotopaxia Mathias & Constance Ref.: Mathias & Constance, 1967. C. asp/undii Mathias & Constance; EC; 3500--4000 m C- whitei Constance & Alverson; CO; 4300 m Daucus L. D. montanus Humb. & Bonpl. ex Spreng.; CR, CO, VE, EC; 1900--4000 m Eryngium L. E. humboldtii F.Delaroche; CO, VE; 2700--4200 m E. humile Cav.; CR, PA, CO, VE, EC, PE; 3100--4300 m; includes-\Íar. brevibracteatum Hieron.·, var. caulescens Kurith, var. pulchellum Wedd., var. radicijlarum (Trattinnick) Hieron. ex. H.Wolff, and var. ste/latum (Mutis) Hieron. Hydracatyle L. H. alchemilloides A.Rich.; EC, PE; 3700--4200 m H. andina Cuatrec.; CO; 3500 m H. banplandii A.Rich. (syn.: H. bonplandii A.Rich. var. · glabra Mathias); CO, VE, EC, PE?; 3300--4100 m H. gunnerifolia Wedd.; CO, VE; 2800--4000 m H. hederacea Mathias; CO, VE; 3700--4000 m H. humbaldtii A.Rich. (syn.: H. humbaldtii A.Rich. var. pubescens Mathias); CO; 3100-3600 m H. incrassata Ruiz & Pav.; CO, PE; 3700-3800 m H. lehmannii Mathias; CO; 3300-3550 m H. mexicana Schltdl. & Cham.; CO; 3300-3600 m H. multifida A.Rich.; CO, VE; 3100-3500 m H. cf. palmata Mathias; PE; 3400-3550 m H. ranunculaides L. f.; CO, VE, PE; 2800-3800 m Lilaeopsis Greene L. mac/aviana (Gand.) A.W.Hill; CO, EC, PE; 3100--4100 m
83
L. schaffneriana (Sch1tdl.) J.M.Coult. & Rose subsp. schaffneriana; CO, EC; 2900--4450 m Micrapleura Lag. M.jlabellifa/ia Mathias; CO;? m [Note: Collected near snow line; and in the uppennost (ecotone) forest (Cieef, pers. comm.)] Myrrhidendran J.M.Coult. & Rose M. chirripaense Suess.; CR; 3300-3900 m M. dannell-smithii J.M.Cou1t. & Rose; CR, PA; 1800-3800 m M. glaucescens (Benth.) J.M.Coult. & Rose; CO, EC; 35004400 m M. maxanii J.M.Coult. & Rose; CR, PA; 3500--4000 m M. pennellii J.M.Coult. & Rose; CO; 3550 m Neanelsania J.M.Coult. & Rose N. acuminata (Benth.) J.M.Coult. & Rose ex Drude; CO; 2100-3400 m Niphagetan Schltdl. Ref.: Mathias & Constance, 1951, 1962, 1967, 1976a. N. azarellaides Mathias & Constance; EC, PE; 3600--4100 m N. chirripoi (Suess.) Mathias & Constance [syn.: N. angustisecta (H.Wo1ff) Mathias & Constance]; CR, PA, CO, VE; 2700-3800 m N. cleefii Mathias & Constance; CO; 3600--4000 m N. colombiana Mathias & Constance; CO; 3250-3500 m N. dissecta (Benth.) J.F.Macbr.; CO, VE, EC, PE; 32004500 m; includes var. ciliata (Wedd.) J.F.Macbr. & var. dissecta N. fruti casa Mathias & Constance; CO; 3750--4050 m N. g/aucescens (Kunth) J.F.Macbr.; CO, EC; 3000--4050 m N.jasei Mathias & Constance; CO; 3000--4500 m N. kalbreyeri (H.Wolff) Mathias & Constance; CO, VE; 3100--4200 m N. /ingula (Wedd.) Mathias & Constance; CR, PA, CO; 3200--4300 m N. pusilla (Wedd.) Mathias & Constance; CO; 3600--4150 m N. sprucei (H.Wolff) Mathias & Constance; EC;? m N. terna/a (Willd. ex Schltdl.) Mathias & Constance; CO, VE, EC; 2800--4000 m; includes var. orientalis Mathias & Constance & var. terna/a Oreamyrrhis Endl. O. andicola (Kunth) Hook.f.; CR, CO, EC, PE; 3100--4500 m Ottaa Kunth O. aenanthoides Kunth; CR, CO, VE, EC; 3100--4300 m; includes var. majar Wedd. & var. aenanthaides Perissacaeleum Mathias & Constance (syn.: Prianasciadium S.Watson) Ref.: Mathias & Constance, 1967. P. barclayiae Mathias & Constance; CO; 3150-3400 m P. crinaideum (Mathias & Constance) Mathias & Constance (syn.: Prianosciadium crinaideum Mathias & Constance); CO; 3250-3850 m
84
MEMOIRS OF THE NEW YORK BOTANICAL OARDEN
P. phylloideum (Mathias & Constance) Mathias & Constan ce . (syn.: Prionosciadium phylloideum Mathias & Constance); CO; 3000-3100 m P. purdiei Mathias & Constance; CO; 3250-4500 m Prionosciadium S.Watson (see Perissocoeleum)
AQUIFOLIACEAE Ilex L. l. colombiana Cuatrec.; CO; 3450-3800 m l. ericoides Loes.; PE; 3500-3650 m l. kunthiana Triana; CO, VE; 3100-3700 m l. myricoides Kunth var. polyphylla (Benth.) Loes.; CO; 2800-3250 m l. pernervata Cua_trec.: CO; 3500-3600 m l. sessiliflora Triana; CO; 3000 m l. spinulosa Cuatrec.; CO; 3500-3600 m J. suprema Cuatrec.; CO; 3100-3750 m J. truxillensis Turcz. var. bullatissima Cuatrec.; VE; 3100_:_ 3400m J. unijlora Benth. var. paramensis Cuatrec.; CO; 3400-3550 m
ARALIACEAE Oreopanax Decne. & P1anch. [Note: The genus is native to montane forest, but the species listed may be found at the upper edges of the forest or in disturbed subpáramo shrubberies.] O. argentatus (Kunth) Decne. & Planch.; CO, EC; 3250 m O. bogotensis Cuatrec.; CO; 2800-4000 m O. chrysoleucus Planch. & Lind.; VE; 3400-4000 m O. crassinervius (Kunth) Decne. & Planch.; CO; 2700-3900 m O. crataegodorus Harms; CO; 3850-3900 m O. discolor (Kunth) Decne. & Planch.; CO; 3000-3500 m O. farallonensis Cuatrec.; CO; 2700-3600 m O.fontquerianus Cuatrec.; CO; 3250-3300 m O. incisus (Willd.) Decne. & Planch.; CO; 3800-4200 m O. killipii Harms; CO; 3800-4200 m O. mutisianus (Kunth) Decne. & Planch.; CO; 3000-3800 m O. nigrus Cuatrec.; CO; 2750-3250 m O. obscurus Borchs.; EC; 2800-3500 m O. seemannianus Marcha! ("seemannianum") [syn.: O. nitidus Cuatrec. ("nitidum")]; CO, EC?; 2400-4000 m O. tolimanus Harms; CO; 3650 m Sche.fflera J.R.Forst. & G.Forst. [Note: A forest genus of mainly hemi-epiphytes, but sorne species are found in subpáramo shrubberies or forest patches.] S. bogotensis Cuatrec.; CO; 2800-4000 m S. manus-dei Cuatrec.; CO; 3500-3600 m S. marginata Cuatrec.; CO; 2800 m S. velutina Cuatrec.; CO; 2800 m
ASCLEPIADACEAE Cynanchum L. C. bricenoi Morillo; VE; 3100-3350 m
[VOL. 84
C. diazmirandae Morillo; VE; 3400-4100 m C. granatense (Baill.) Morillo; CO; ? m C. intricatum K.Schum.; EC; 3900-4200 m C. longecalicinum Morillo; EC; 3700-3800 m C. paramorum Morillo; VE; 3000-3200 m C. rugosum (Turcz.) Morillo (syn.: Metastelma rugosa Turcz.); CO, VE; 2925-3500 m
Metastelma R.Br. (see Cynanchum) Sarcostemma R.Br. S. lysimachioides (Wedd.) R.W.Holm; PE; 3000-3750 m
ASTERACEAE ·'Ref.: Anderberg, 1991; Aristeguieta, 1964; Badillo, 1994, 1997; Bremer, 1994; Cuatrecasas, 1969, 1976, 1985b; Díaz Piedrahita & Vélez-Nauer, 1993; Dillon & Sagástegui, 1991a, 1991b; Jeffrey, 1992; King & Robinson, 1987; McVaugh, 1984; Nesom, 1994b, 1994c; Sagástegui & Dillon, 1986; Vision & Dillon, 1996; Weddell, 1855-1857. Achillea L. [Note: lntroduced from Europe and now escaped.] A. millefolium L.; CR, CO, EC; 2800-3500 m Achyrocline (Less.) DC. A. afata (Kunth) DC. (syn.: Gnaphalium rufescens DC., G. pellitum Kunth); CO, EC, PE; 3300-4100 m A. bogotense (Kunth) DC.; CO, VE; 3400-4000 m A. celosioides (Kunth) DC.; PE; 2200-3300 m A. crassipes S.F.Biake; CO; 3150-3700 m A. hallii Hieron.; EC; 2800-3300 m A. lehmannii Hieron.; CO, VE; 3150-4300 m A. mollis Benth.; CO; 3100-3300 m A. moritziana Klatt [syn.: Gnaphalium moritziana (Kiatt) Aristeg.; Pseudognaphalium moritzianum (Kiatt) V.M.Badillo]; CO, VE, EC; 3100-3930 m A. satureioides (Lam.) DC. [syn.: A. candicans (Kunth) DC.]; CO, VE, EC, PE; 3400-4000 m A. trianae Klatt; CO, EC; 3000-3700 m Aequatorium B.Nord. A. albiflorum (Wedd.) Cuatrec. & S.Díaz (syn.: Gynoxys albiflora Wedd.); CO; 2000-3900 m A. latibracteolatum S.Díaz & Cuatrec.; CO; 3300-3350 m A. polygonoides (Cuatrec.) B.Nord. (syn.: Senecio polygonoides Cuatrec.); CO; 3400-3700 m A. verrucosum (Wedd.) S.Díaz & Cuatrec. (syn.: Gynoxys verrucosa Wedd. var. verrucosa and var. grandifolia Cuatrec.); CO; 3200-3900 m Aetheolaena Cass. (see Lasiocephalus) Ageratina Spach A. ampla (Benth.) R.M.King & H.Rob.; CO; 3400-4000 m A. angustifolia (Kunth) R.M.King & H.Rob.; CO; 20503575 m A. anisochroma (Klatt) R.M.King & H.Rob.; CR; 2000-3600 m
.{
1999]
PARAMOS
A. aristeguietii R.M.King & H.Rob.; VE; 3200--4025 m A. articu/ata (Sch.Bip. ex Hieran.) R.M.King & H.Rob.; CO, VE, EC; 2800--4000 m A. asclepiadea (L.f) R.M.King & H.Rob.; CO; 2800-3100 m A. azangaroense (Sch.Bip. ex Wedd.) R.M.King & H.Rob.; CO, EC, PE; 1700--4150 m A. baccharoides (KuntlÚ R.M.King & H.Rob.; CO; 28004000 m A. chachapoyensis (Cuatrec.) R.M.King & H.Rob.; PE; 3100-3550 m A. chiriquensis (B.L.Rob.) R.M.King & H.Rob.; PA; 3350-3400 m A. crassiceps (B.L.Rob.) R.M.King & H.Rob.; CO; 38004100m A. cuatrecasii R.M.King & H.Rob.; CO; 2800-3100.m A. cuencana (B.L.Rob.) R.M.King & H.Rob.; EC; 35003700 m A. cutervensis (Hieran.) R.M.King & H.Rob.; EC; 32503700 m A. e/egans (Kunth) R.M.King & H.Rob.; CO; 2800--4000 m A. exertovenosa (Klatt) R.M.King & H.Rob.; EC, PE; 32003500 m A.fastigiata (Kunth) R.M.King & H.Rob.; CO; 3000-3700 m A.flaviseta (B.L.Rob.) R.M.King & H.Rob.; CO; 2600-3800 m A. g/yptoph/ebia (B.L.Rob.) R.M.King & H.Rob.; CO, EC; 3250--4050 m A. graci/is (Kunth) R.M.King & H.Rob. var. epilobioides (Kunth) R.M.King & H.Rob.; CO, VE; 1700--4050 m A. gynoxoides (Wedd.) R.M.King & H.Rob.; CO; 3350--4000 m A. ixiocladon (Benth.) R.M.King & H.Rob.; CR, PA; 10003800m A.jahnii (.EÍ.L.Rob.) R.M.King & H.Rob. [syn.: A. stevioides (Steyerm.) R.M.King & H.Rob.]; VE; 2600--4000 m A. kupperi (Suess.) R.M.King & H.Rob.; CR, PA; 34003700m A. la lipes (Benth.) R.M.King & H.Rob.; CO; 2800-3500 m A. mutiscuensis (B.L.Rob.) R.M.King & H.Rob.; CO; 28804200 m A. ocanensis (B.L.Rob.) R.M.King & H.Rob. (syn.: Eupatorium ocanense B.L.Rob.); CO; 3200-3700 m A. paramense (Aristeg.) R.M.King & H.Rob.; VE; 27003200m A. pichinchensis (Kunth) R.M.King & H.Rob. [syn.: A. ibaguensis (Sch.Bip. ex Hieran.) R.M.King & H.Rob.]; CO, VE, EC; 3000--4500 m A. prunifolia (H.B.K.) R.M.King & H.Rob.; EC; 2800-3400 m A. pseudochilca (Benth.) R.M.King & H.Rob. (syn.: Eupatorium cotacachense Hieran.); CO, EC; 2600-3500 m A. scopulontm (Wedd.) R.M.King & H.Rob.; PE; 3000-3800 m A. sodiroi (Hieran.) R.M.King & H.Rob.; EC; 2500-3400 m A. sternbergiana (DC.) R.M.King & H.Rob.; PE; 2800-3450 m A. theaefolia (Benth.) R.M.King & H.Rob.; CO, VE; 34004100 m; including var. subel/iptica B.L.Rob. & var.
theaefolia A, tinifolia (Kunth) R.M.King & H.Rob.; CO, VE; 3000-
Alloispermum Willd. Ref.: Fernández, 1980.
A. caracasanum (Kunth) H.Rob. [syn.: Catea caracasana (Kunth) Kuntze]; CO, VE; 3300--4000 m
A. insuetum Fernández, Urbatsch & Sullivan; CO; 24503050 m
A. sodiroi (Hieran.) H.Rob.; CO; 2650-3000 m Antennaria Gaertn. A. linearifo/ia Wedd. [syn.: Gnaphalium linearifolium (Wedd.) Franchet]; EC, PE; 3100--4000 m
Aphanactis Wedd. Ref: Robinson, 1997; Turner, 1980.
A. antisanensis H.Rob.; EC; 4200--4400 m A. barclayae H.Rob.; EC; 4100--4300 m A. brevi/igulata (see Se/loa breviligulata) A. cocuyensis Cuatrec.; CO; 4100 m A.jamesoniana Wedd.; CO, EC; 3500--4300 m A. ligulata (see Sel/oa ligulata) A. oel/gaardii H.Rob.; EC; 3500--4600 m A. pi/ose/loides Cuatrec.; CO; 3100--4300 m A. vi/losa S.F.Blake; EC, PE; 3500--4200 m Aristeguietia R.M.King & H.Rob. Ref.: King & Robinson, 1975.
A. anisodonta (B.L.Rob.) R.M.King & H.Rob.; PE; 22003450 m
A. arborea (Kunth) R.M.King & H.Rob. (syn.: Eupatorium arboreum Kunth); EC; 3180-3500 m A. buddleaifolia (Benth.) R.M.King & H.Rob. (syn.: Eupatorium buddleaifolium Benth.); PE; 3300 m A. caca/oides (Kunth) R.M.King & H.Rob.; EC; 3100-3350 m A. glutinosa (Lam.) R.M.King & H.Rob. (syn.: Eupatorium glutinosum Lam.); CO, EC; 3500-3950 m Ascidiogyne Cuatrec. A. sanchezvegae Cabrera; PE; 3600--4500 m A. wurdackii Cuatrec.; PE; 3200-3900 m Aster L. (see Noticastrum) Baccharis L.
B. B. B. B. B. B. B. B.
3700 m
A. vaccinifolia (Benth.) R.M.King & H.Rob.; CO; 32503600m
A. vallincola (DC.) R.M.King & H.Rob.; PE; 2800-3500 m
85
B. B.
[Note: Species of this genus are found mostly in subpáramo shrubberies.] Ref: Cuatrecasas, 1967a, 1967b, 1968, 1969. alaternoides Kunth; EC; 3300-3900 m arbutifolia (Lam.) Vahl; EC; 3600--4600 m; includes var. arbutifolia and var. jamesonii Cuatrec. aretioides Turcz.; EC; 4300 m auriculigera Hieran.; PE; 2700-3500 m barragensis Cuatrec.; CO; 3600-3700 m bogotensis Kunth; CO; 2800--4100 m boyacensis Cuatrec.; CO; 3200-3400 m brachy/aenoides DC.; CO, VE, EC; 2800-3450 m; ineludes var. brachylaenoides and var. oblanceolata (Hieran.) Cuatrec. buddleioides Kunth; CO, EC; 2590-3420 m buxifo/ia (Lam.) Pers.; CO, EC; 3300--4200 m
86
MEMOIRS OF THE NEW YORK BOT ANICAL O ARDEN
B. taespitosa (Ruiz & Pav.) Pers. (syn.: B. alpina Kunth); CO, EC, PE; 3600--4700 m; includes var. alpina (Kunth) Cuatrec., var. alpicola Cuatrec., and var. caespitosa B. caldasiana Cuatrec.; CO; 3400--4300 m B. cochensis Hieran.; CO, EC; 2850-3300 m B. cutervensís Hieran.; EC, PE; 2400-3400 m B. emarginata (Ruiz & Pav.) Pers.; PE; 2400-3300 m B.fi'igida Kunth; PE; 3100 m B. genístelloides .(Lam.) Pers.; CO, EC, PE; 3300--4300 m B. granadina Cuatrec.; CO, EC; 3200 m B. grandicapitulata Hieran. var. grandícapítulata; PE; 25004000m B. grandiflora Kunth; CO, EC; 2800-4000 m; includes subsp.faralloensís Cuatrec. and subsp. grandzjlora B. huairacajensis Hieran.; E¡C; 3000-3500 m B. humifitsa Kunth; EC; 3800--4200 m B. hutchínsonii Cuatrec.; PE; 3000-3SOO m B. klattií Benaist; EC; 3400--4300 m B. latifolia (Ruiz & Pav.) Pers. (syn.: B.jloribunda Kunth); CO, VE, EC, PE; 3200--4000 m B. ledifolia Kunth; PE; 3200-3500 m B. lehmannii Klatt; CO; 3000-3550 m B. libertadensís (S.B.Janes) H.Rab.; PE; 2900-3400 m B. macrantha Kunth (syn.: B. cundínamarcensis Cuatrec.); CO, EC; 3200-4000 m; includes subsp. caucacensis Cuatrec., subsp. cundinamarcensis (Cuatrec.) Cuatrec. and its var. cocuyensis Cuatrec., subsp. macrantha, and subsp. minar (Cuatrec.) Cuatrec. B. marcetíifolia Benth.; CO, EC; 3200--4000 m B. oblongifolia (Ruiz & Pav.) Pers.; EC; 2400-3400 m B. odorata Kunth; CO, EC; 2200--4000 m B. pachycepha/a Hieran.; PE; 2700-3700 m B. padifolia Hieran.; CO, EC; 3500--4250 m B. paramíco/a Cuatrec.; CO; 3300--4000 m B. peruviana Cuatrec.; PE; 3000-3900 m B. prunifolia Kunth (syn.: B. densifolia Rus by); CO, EC, VE; 3300-4500 m; includes var. prunifo/ia, var. sierrana Cuatrec., and var. subprunifolia Cuatrec. B. revo/uta Kunth; CO; 3100--4000 m B. rupíco/a Kunth (syn.: B. capitoides Cuatrec.); CO; 31004350 m; includes var. orienta lis Cuatrec., var. rupicola, var. serrata Cuatrec., and var. summapacis Cuatrec. B. tarmensis Cuatrec.; PE; 2600-3600 m B. teindalensis Kunth; CO, EC; 2800-3700 m B. tricuneata (L.f.) Pers. (syn.: B. microphylla Kunth); CO, VE, EC, PE; 2800--4400 m; includes var. antioquensis Cuatrec., var. paramontm Cuatrec., var. parvifolia Cuatrec., var. procumbens Cuatrec., var. tolimensis (Hieran.) Cuatrec., and var. trícuneata B. uniflora (Ruiz & Pav.) Pers.; PE; 3300--4000 m B. vacciniifolia Cuatrec.; CO; 3200-3700 m B. venosa (Ruiz & Pav.) Pers.; PE; 3500-3750 m Badilloa R.M.King & H.Rab. B. salicina (Lam.) R.M.King & H.Rab.; EC; 3700-3750 m B. venezuelense (V.M.Badilla) R.lVÍ.King & H.Rab.; VE; 2200-2900m
[VOL. 84
Barnadesia Mutis ex L. f. Ref.: Chung, 1965. B. arborea Kunth (syn.: B. sodiroi Hieran.); EC; 3100--4250 m B. dombeyana Less.; PE; 2700--4000 m B. spínosa L.f.; CO, EC; 2600--4100 m Be/loa Remy B. longifolia (Cuatrec. & Aristeg.) Sagást. $<- M.O.Dillan (syn.: Lucilia /ongifolia Cuatrec. & Aristeg.); CO, VE, EC, PE; 3600--4300 m B. lopezmírandae Cabrera; PE; 3100 m B. piptolepis (Wedd.) Cabrera (syn.: Lucilia pípto/epís Wedd.); CO, VE, EC, PE; 2350--4300 m B.plícatifolia Sagást. & M.O.Dillan; PE; 3000-3500 m B.'radíans (Benth.) Sagást. & M.O.Dillan [syn.: Lucilia radí. ans (Benth.) Cuatrec.]; CO, VE, EC, PE; 3100--4300 m B. spathulifolia Sagást. & M.O.Dillan; PE; 3200-3800 m B. turno·¡' Sagást. & M.O.Dillan; PE; 3200--4000 m Bídens L. Ref.: Sherff, 1937. B. andico/a Kunth; CO, VE, EC, PE; 2900--4300 m B. humilis Kunth; CO, VE, EC; 3000--4200 m B. obstruthoídes (DC.) Sch.Bip.; CR; 2000-3500 m B. pusil/a Sherff; EC; 3700 m B. rubifolia Kunth; CO, VE; 1800-3700 m B. tríplinervia Kunth; CO, VE, EC, PE; 3100--4000 m; ineludes var. boyacana Sherff, var. macrantha (Wedd.) Sherff, and var. mollis (Paepp. & Endl.) Sherff B/akiella Cuatrec. B. bartsíifolia (S.F.Blake) Cuatrec.; CO, VE; 3400--4200 m Cabreríel/a Cuatrec. C. opposítícordís (Cuatrec.) Cuatrec. (syn.: Senecio oppositícordius Cuatrec.); CO; 2970-3000 m C. sanctae-martae (Greenm.) Cuatrec.; CO; 2400-3200 m Calea L. (see Sabazía and Alloispermum) Chamomilla Gray (see Matricaria) Chaptalia Vent. C. anisobasis S.F.Blake; CO; 3200-3850 m C. cordata Hieran.; CO, EC, PE; 3500--4200 m C. e f. incana Cuatrec.; CO; 3500--4050 m C. merídensís S.F.Blake; VE; 2000-3600 m C. nutans (L.) Palak; VE; 3400-3600 m C. paramensís Cuatrec.; CO, VE; 3250-3350 m Chersodoma Phi!. Ref.: Dillan & Sagástegui, 1996. C. deltoídea M.O.Dillan & Sagást.; PE; 3450--4000 m Chevreulia Cass. C. acumínata Less.; PE; 1600-3500 rri
1999]
PARAMOS
Chionolaena DC. Ref.: Freire, 1993. C. cf. ch1ysocoma (Wedd.) Freire; CO; 3300-4300 m C. columbiana S.F.Blake; CO; 3250-4450 m Chromolaena DC. C." bu/lata (Klatt) R.M.King & R.Rob.; CO; 2625-3575 m [Note: This species may be equal to C. hypericifolia.] C. colombiana (Reering) R.M.King & R.Rob.; CO; 34503750m C. hypericifolia (Kunth) R.M.King & H.Rob.; CO; 700-3700 m C. persericea R.M.King & R.Rob.; CO, VE; 2800-3100 m Chrysactinium (Kunth) Wedd. Ref.: Zermoglio & Funk, 1997. C. acaule (Kunth) Wedd. [syn.: C. longiradiatum (Rieron.) R.Rob. & Brettell]; EC, PE; 2800-4300 m C. breviscapum Sagรกst. & M.O.Dillon; PE; 3300-3600 m C. caulescens (Rieron.) R.Rob. & Brettell; EC, PE; 3200 m C. hieracioides (Kunth) H.Rob. & Brettell; EC, PE; 30004300m C. rosulatum (Rieron.) R.Rob. & Brettell; PE; 3450-4000 m Chuquiraga Juss. Ref.: Ezcurra, 1985. C. jussieui J.F.Gme!. [syn.: C. insignis (Willd.) Rumb. & Bonpl., C. lancifolia Humb. & Bonpl.]; CO, EC, PE; 3400-4750 m C. oblongifolia Sagรกst. & Sรกnchez Vega; PE; 3300 m C. weberbaueri Tovar; PE; 3000-4400 m Cirsium Mili. [Note: The genus and species are introduced European weeds in pรกramo.] C. subcoriaceum (Less.) Sch.Bip.; CR; 1900-3500 m C. vulgare (Savi) Ten.; EC; 3400-3500 m Coespeletia Cuatrec. C. albarregensis Cuatrec., sp. nov. ined.; VE; 3850 m [Note: This species is not meant to be published herein.] C. elongata (A.C.Sm.) Cuatrec. (syn.: Espeletia elongata A.C.Sm.); VE; 3300-3900 m C. moritziana (Sch.Bip.) Cuatrec. (syn.: Espeletia moritziana Sch.Bip.); VE; 3700-4400 m C. spicata (Sch.Bip. ex Wedd.) Cuatrec. [syn.: Espeletia alba A.C.Sm., E. spicata Sch.Bip. ex Wedd.; C. alba (A.C.Sin.) Cuatrec.]; VE; 3000-4500 m C. thyrsiformis (A.C.Sm.) Cuatrec. (syn.: Espeletia marcan a Cuatrec., E. racemosa Cuatrec., E. thyrsiformis A.C.Sm.); VE; 2800-3400 m C. timotensis (Cuatrec.) }:uatrec. [syn.: Espeletia timotensis Cuatrec., E. lutescens Cuatrec. & Aristeg.; C. lutescens (Cuatrec. & Aristeg.) Cuatrec.]; VE; 3900-4500 m Conyza Less. Ref.: Cuatrecasas, 1969. C. bonariensis (L.) Cronquist var. leiotheca (S.F.Biake)
87
Cuatrec. [syn.:. C. jloribunda Kunth, C. sumatrensis (Retz.) E.Walker]; CO, VE, EC; 3000-3400 m C. cardaminifolia Kunth; EC; 3450-4600 m C. coronopifolia Kunth; CO, EC, PE; 2800-4000 m C.filaginoides (DC.) Rieron. (syn.: Laenneciafilaginoides DC.); CO, VE; 3200 m C. gnaphalioides Kunth [syn.: Laennecia gnaphalioides (Kunth) Cass.]; CO, VE, EC, PE; 3200 m C. laevigata (Rich.) Pruski (syn.: C. apurensis Kunth); VE; 3300-3350 m C. lasseriana Aristeg. [syn.: Laennecia lasseriana (Aristeg.) G.L.Nesom]; VE; 2500-4100 m C. mima S.F.Biake [syn.: Laennecia mima (S.F.Blake) G.L.Nesom]; VE; 3500-4300 m C. primuliifolia (Lam.) Cuatrec. & Lourteig (syn.: C. chilensis Spreng.); EC; 3600-4000 m C. prolialba Cuatrec.; CO, EC; 3350-4300 m C. sophiifolia Kunth (syn.: Laennecia sophiifolia (Kunth) G.L.Nesom]; CO, VE, EC; 2100-3700 m C. trihecatactis (S.F.Biake) Cuatrec. [syn.: Erigeron trihecatactis S.F.Biake]; CO, EC; 3000-3900 m C. uliginosa (Benth.) Cuatrec. [syn.: Erigeron popayanensis Rieron.); CO, VE, EC; 3100-4500 m; includes var. angustifolia Cuatrec., var. columbiana (Rieron.) Cuatrec., var. hirsuta (Rieron.) Cuatrec., and var. uliginosa] Coreopsis L. Ref.: Tadesse et al., 1995. C. fasciculata Wedd. (syn.: C. suaveolens Sherff var. ecuadoriensis Sherff); EC, PE; 2700-4000 m C. integra S.F.Biake; PE; 2250-3750 m C. obovatifolia Sagรกst.; PE; 2700-3600 m C. petrophiloides B.L.Rob. & Greenm.; CR; 3300 m C. polyactis S.F.Biake & Sherff ex Sherff; PE; 3500-3600 m C. venusta Kunth (syn.: C. !angula S.F.Biake); EC, PE; 2100-4100 m C. woytkowskii Sherff; PE; 2700-3400 m Cotufa L. C. australis (Sieber ex Spreng) Roo k. f.; PE; 3100-3200 m C. mexicana (DC.) Cabrera [syn.: C. minuta (L. f.) Schinz]; CR, CO, VE, EC, PE; 3000-4400 m Critoniopsis Sch.Bip. C. huairacajana (Rieron.) H.Rob. (syn.: Vernonia huairacajana Rieron.); EC, PE; 2900-3450 m Cronquistianthus R.M.King & R.Rob. C. celendensis R.M.King & R.Rob. [syn.: Eupatorium celendensis (R.M.King & R.Rob.) B.L.Turner]; PE; 2100-3100m C. desmophyllus (B.L.Rob.) R.M.King & R.Rob. (syn.: Eupatorium desmophyllus B.L.Rob.); PE; 3100-3200 m C. lavandulaefolia (DC.) R.M.King & H.Rob. (syn.: Eupatorium lavandulaefolium DC.); PE; 3200-3400 m C. pseudoriganoides (Rieron.) R.M.King & R.Rob. (syn.: Eupatorium pseudoriganoides Rieron.); EC; 3200-3700 m
1
88
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
Ct<afrecasasie/la H.Rob. C. isernii (Cuatrec.) H.Rob. (syn.: Luci/iopsis isernii Cuatrec.); EC; 3700-4700 m Cu/citium Bonpl. (see also synonyms under Lasiocephalus and Senecio) Dendrophorbium (Cuatrec.) C.Jeffrey (see also synonyms under Pentacalia and Senecio) Diplostephium Kunth Ref.: Cuatrecasas, 1943, 1969. D. abietinum S.F.B1ake; PE; 4000 m D. alveo/atum Cuatrec.; CO; 3400-4450 m D. anactinotum Wedd.; CO; 3600-4430 m D. apiculatum S:F.Blake; CO; 3500-4200 m D. asp/undii Cuatrec.; EC; 3500 m D. azureum Cuatrec.; PE; 4000 m D. barclayanum Cuatrec.; EC; 3700 m D. bicolor S.F.Blake (syn.: D. tabanense Cuatrec.); CO; 3350-3700 m D. cajamarquillensis Cuatrec.; PE; 3925 m D. ca/laense Cuatrec.; PE; 3300-4000 m D. cayambense Cuatrec.; CO, EC; 3250-3800 m D. cinerascens Cuatrec. subsp. puracense (Cuatrec.) Cuatrec. (syn.: D. vio/aceum Cuatrec. var. puracensis Cuatrec., D. cinerascens var. centra/e Cuatrec.); CO, EC; 32004200 m D. co/ombianum (Cuatrec.) Cuatrec.; CO; 3700-4500 m D. coriaceum Cuatrec.; CO; 3100-3400 m D. costaricense S.F.Blake; CR, PA; 3300-3800 m D. crassifolium Cuatrec.; CO, VE; 3000-3600 m D. crypteriophyllum Cuatrec.; EC; 3000-3400 m D. cyparissias Wedd.; CO; 3500-4000 m D. empetrifolium S.F.Blake; EC; 2800-3400 m D. ericoides (Lam.) Cabrera (syn.: D. antisanense Hieron., D. lavandu/ifolius Kunth, D. pycnophyllum S.F.Blake); EC; 3000-4500 m D. eriophorum Wedd.; CO; 3300-4600 m D. espinosae Cuatrec.; EC; 2700-3800 m D.farallonense (Cuatrec.) Cuatrec.; CO; 3300-4000 m D..fernandez-a/onsoi S.Díaz; CO; 3250-3300 m D. jloribundum (Benth.) Wedd.; CO, EC; 3400-3700 m; includes subsp. aequatoriense Cuatrec., s~bsp. cundinamarcense Cuatrec., subsp. jloribundum, and subsp. putumayense Cuatrec. D.foliosissimum S.F.Blake·; EC, PE; 2900-4000 m D. friedbergii Cuatrec.; PE; 3250-3400 m D.ji·ontinense Cuatrec.; CO; 3200-3450 m D. glandu/osum Hieron. (syn.: D. cicatricosu.m S.F.Biake); CO, EC; 3400-4100 m D. glutinosum S.F.Blake (syn.: D. subspathu/atum Cuatrec.); CO, VE; 3700-4200 m; includes f. glutinosum, f. microphyllum Cuatrec., and f. subspathu/atum (Cuatrec.) Cuatrec. D. grantii Cuatrec.; CO; 2550-? m D. hartwegii Hieron. (syn.: D. adenachaenium S.F.Blake); CO, EC; 3250-4200 m D. heterophyllum Cuatrec.; CO; 3400-3800 m
[VOL. 84
D. inesianum Cuatrec.; CO; 3500-4500 m D.je/skii Hieron.; EC; 3350-3450 m D. juajibioyi Cuatrec.; CO; 3550-3900 m; includes subsp. juajibioyi and subsp. leucopappum Cuatrec. D. /acunosum Cuatrec.; CO; 3800-4300 m D. /eiocladum S.F.Blake; CO; 3300-3500 m D. macrocephalum S.F.B1ake; EC; 3200-3800 m D. meyenii (Sch.Bip. ex Wedd.) S.F.Blake; PE; 3200-3700 m D. micradenium S.F.Blake; CO; 3300-3500 m D. nevadense Cuatrec.; CO; 3300-4700 m D. oblanceolatum S.F.Blake; EC; 3000-3950 m D. ob/ongifolium Cuatrec.; CO; 2770 m D. obtusum S.F.Blake; VE; ~200-3400 m D. ochraceum (Kunth) N e es (syn.: D. denticulatum .· S.F.Blake); CO, EC; 3000-3500 m D. pachyphyllum Cuatrec.; PE; 3300-3600 m D. parvifolium S.F.Blake (syn.: D. microphyllum Wedd.); CO; 3000-4500 m D. phylicoides (Kunth) Wedd.; CO; 2800-4050 m D. pittieri Cuatrec.; CO; 3300-3600 m D. rangelii Cuatrec.; CO; 3850 m D. revolutum S.F.Blake (syn.: D. revolutum var. rubrum Cuatrec., D. revolutum f. macrocephala Cuatrec.); CO; 3000-4200 m D. rhododendroides Hieron.; CO, EC; 3500-4000 m D. rhomboidale Cuatrec. [syn.: D. cochense Cuatrec., D. lehmannianum (Hieron.) Hieron.]; CO, EC; 3400-4400 m; includes var. paucijlorum Cuatrec. and var. rhomboidale D. ritterbushii Cuatrec.; CO; 4200-4300 m D. rosmarinifolium (Benth.) Wedd. (syn.: D. baccharideum S.F.Blake); CO, VE; 2500-3300 m D. rupestre (Kunth) Wedd.; CO, EC; 3600-4500 m D. sagasteguii Cuatrec.; PE; 3800-4000 m D. santamartae Cuatrec.; CO; 3450 m D. saxati/e Cuatrec.; CO; 3850-4050 m D. schultzii Wedd. (syn.: D. pleistogynum S.F.Blake); CO, VE, EC; 3100-4400 m; includes var. lehmannianum Hieron., var. orienta/e (Cuatrec.) Cuatrec., and var. sch'ultzii D. spinulosum Wedd.; CO, EC; 3300-4000 m D. stuebelii (Hieron.) Cuatrec. (syn.: Aster stuebelii Hieron.); PE; 3200-3450 m (Note: Perhaps this species is better placed in Noticastrum.] D. tenuifolium Cuatrec.; CO; 2600-3900 rn D. tergocanum Cuatrec.; CO; 3750-3800 111 D. tolimense Cuatrec. (syn.: D. boyacense Cuatrec.); CO; 3200-4000 m D. venezue/ense Cuatrec.; VE; 3300-3800 m D. weddellii S.F.Blake (syn.: D. sessiliflorum Wedd.); CO; 3400-4400 m D. wurdackii Cuatrec.; PE; 3200-3450 m D. yahuarcochense Cuatrec.; PE; 3600 m Dorobaea Cass. Ref.: Nordenstam & Pruski, 1995. D. ca/lacal/ensis (Cllatrec.) B.Nord. & Pruski (syn.: Senecio ca/laca/lensis Cuatrec.); PE; 2800-3100 m
1999]
PARAMOS
D. pimpinelifolia (Kunth) B.Nord. (syn.: Senecio pimpinelifolius Kunth, S. nubigenus Kunth); CO, EC, PE; 34004200 m
89
E. estanis/ana Cua(rec.; CO; 3400-4100 m E.ji-ontinoensis Cuatrec.; CO; 3300-3500 m E. grandiflora Humb. & Bonpl. (syn.: E. grandiflora var. subnivalis Cuatrec.); CO; 3200-4100 m; inc1udes var. Erígeron L. attenuata Cuatrec., var. boyacana Cuatrec., and var. Ref.: Solbrig, 1962. grandiflora E. guascensis Cuatrec.; CO; 3200-3300 m [Note: This speE. chionophilus Wedd. var. chionophilus; CO, EC, PE; 3500-4700 m cies may be a hybrid, according to Cuatrecasas E. ecuadoriensis Hieron: (syn.: E. weddellianus Hieron.); (unpubl.).] E. hartwegiana Cuatrec.; CO; 3100-4500 m; includes subsp. CO, EC, PE; 3600-4600 m • E. gnaphalioides Kunth; EC; 3500 m barragansis Cuatrec., subsp. centroandina Cuatrec., and E. irazuensis Greenm.; CR; 1500-3500 m [Note: This spesubsp._hartwegiana var. hartwegiana, var. morarum cies may equa1 E. karwinskianus DC. (J. Sánchez G., Cuatrec., and var. vegasana Cuatrec. pers. comm.).] E. idroboi Cuatrec.; CO; 3100 m E. lanceolatus Wedd. var. subacaulis Wedd.; PE; 2900-3150 m E. incana Cuatrec.; CO; 2900-3900 m E. paramensis Aristeg. & Cuatrec. [syn.: Oritrophium E.jajoensis Aristeg. [syn.: Espeletiopsisjajoensis (Aristeg.) paramense (Aristeg. & Cuatrec.) Aristeg.]; CO, VE; Cuatrec.]; VE; 2600-3600 m 3500-4500 m E.jaramilloi S.Díaz; CO; 3500-3750 m E. raphaelis Cuatrec.; CO, VE; 3100-3650 m E. killipii Cuatrec.; CO; 3100-4000 m; inc1udes var. E. turczaninowii Wedd.; EC; 3100-4400 m (E. pi~natus chisacana Cuatrec. and var. killipii Turcz. auct., nom. illeg.) E. lopezii Cuatrec.; CO; 3100-4500 m; in eludes fo. alticola E. uliginosum Benth.; EC; 3400-3500 m Cuatrec., var. escobalensis Cuatrec., var. /opezii, and var. • E. variifolius S.F.B1ake; EC; 3500-4500 m majar Cuatrec. E. marthae Cuatrec.; VE; 3000-3500 m Espeletia Mutis ex Humb. & Bonpl. (see a1so synonyms E. miradorensis Cuatrec. (syn.: E. grandiflora Humb. & · ·under Coespeletia, Espeletiopsis, Libanothamnus, and Bonpl. var. miradorensis Cuatrec.; CO; 3550 m Ruilopezia) E. murilloi Cuatrec.; CO; 2700-3600 m Ref.: Smith & Koch, 1935. E. nana Cuatrec.; VE; 3000-4000 m E. algodonosa Aristeg.; VE; 3000 m [Note: This species is E. nemenkenii Cuatrec.; CO; 3000-3550 m a hybrid between E. schultzii andE. nana (Cuatrecasas, E. accidenta lis A.C.Sm.; CO; 2600-3000 m; includes subsp. antioquiensis Cuatrec. and subsp. occidentalis unpuql.).] E. annemariana Cuatrec.; CO; 3100-4000 m E. oswaldiana S.Díaz; CO; 2900-3000 m E. perijaensis Cuatrec.; CO, VE; 2700-3300 m E. arbelaezii Cuatrec.; CO; 3300 m E. pescana (S.Díaz) Cuatrec. (syn.: E. brachyaxiantha E. argentea Humb. & Bonpl. [syn.: E. phaneractis S.Díaz subsp. pescana S.Díaz); CO; 3750 m (S.F.B1ake) A.C.Sm.]; CO; 3200-3550 m E. aristeguietana Cuatrec.; VE; 2500-2850 m E. praeji-ontina Cuatrec.; CO; 3450-3500 m E. pycnophy/la Cuatrec.; CO, EC; 3000-4000 m; inc1udes E. aurantia Aristeg.; VE; 4000-4100 m [Note: This speCies subsp. angelensis Cuatrec. and subsp. pycnophy/la var. is a hybrid with Coespeletia sp. (Cuatrecasas, unpubl.).] /acinulata Cuatrec. and var. galerana Cuatrec. E. azucarina Cuatrec.; CO; 4000 m E. robertii Cuatrec.; CO; 3100-3900 m E. barclayana Cuatrec.; CO; 3600-3700 m E. rosita e Cuatrec. (syn.: E. rositae Cuatrec. var. E. batata Cuatrec.; VE; 3600-4300 m macrocephala Cuatrec.); CO; 3300-4050 m E. boyacensis Cuatrec.; CO; 3000-3900 m E. schultesiana Cuatrec.; CO; 2500-3200 m E. brachyaxiantha S.Díaz; CO; 3800-3900 m E. schultzii Wedd.; VE; 2600-4200 m E. brassicoidea Cuatrec.; CO, VE; 2500-3600 m; includes E. semiglobulata Cuatrec. (syn.: E. rufescens Cuatrec.); VE; subsp. angustata Cuatrec., subsp. brassicoidea, and 3400-3900 m subsp. constricta Cuatrec. E. standleyana A.C.Sm.; CO; 2900-3920 m; includes subsp. E. cabrerensis Cuatrec.; CO; 3200 m ampla (Cuatrec.) Cuatrec. and subsp. luxior Cuatrec. _ E. canescens A.C.Sm.; CO; ? m E. steyermarkii Cuatrec.; VE; 2800-3300 m E. cayetana (Cuatrec.) Cuatrec. (syn.: E. grandiflora Humb. E. subnerifolia Cuatrec.; CO; 3700-3950 m & Bonpl. var. cayetana Cuatrec.); CO; 3500-3650 m E. summapacis Cuatrec.; CO; 3650-4000 m E. chocontana Cuatrec.; CO; 3650-3700 m . E. tapirophila Cuatrec.; CO; 3400 m E. cleefii Cuatrec.; CO; 3500-4300 m E. tenore Aristeg.; VE; 3300-3500 m E. congestiflora Cuatrec.; CO; 3150-3900 m E. ti/lettii Cuatrec.; VE; 3100 m E. conglomerata A.C.Sm.; CO; 3400-3900 m E. tunjana Cuatrec.; CO; 2900--3100 m E. cuniculorum Cuatrec.; VE; 33.50 m E. ulotricha Cuatrec.; VE; 3100-3550 m E. curialensis Cuatrec.; CO; 3000-3500 m E. uribei Cuatrec.; CO; 3250-3500 m E. discoidea Cuatrec.; CO; 3200-3800 m E. weddellii Sch.Bip. ex Wedd.; VE; 3100-4000 m E. dugandii Cuatrec.; CO; 3100-3400 m
90
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
Espeletiopsis Cuatrec. Ref.: Cuatrecasas, 1996. E. almorzana Cuatrec.; CO; 3100-3800 m E. angustifolia (Cuatrec.) Cuatrec.; VE; 2600-3500 m E. bogotensis (Cuatrec.) Cuatrec.; CO; 2500-3600 m E. caldasii (Cuatrec.) Cuatrec. (syn.: Espeletia caldasii Cuatrec.); CO; 3200-3400 m E. colombiana (Cuatrec.) Cuatrec . (syn.: Espeletia colombiana Cuatrec.); CO; 4250-4500 m E. corymbosa (Humb. &. Bonpl.) Cuatrec. (syn.: Espeletia corymbosd Humb. & Bonpl.); CO; 3150-4000 m; ineludes subsp. corymbosa and subsp. zipaquirana Cuatrec. E. cristalinensis (Cuatrec.) Cuatrec.; VE; 2400-3100 m E.funckii (Sch.Bip. ex Wedd.) Cuatrec.; CO; 3600-3800 m E. garciae (Cuatrec.) Cuatrec.; CO; 2600-2900 m E. insignis (Cuatrec.) Cuatrec.; CO; 2800-3300 m E.jimenez-quesadae (Cuatrec.) Cuatrec.; CO; 3500-3850 m; includes var. guacharaca (S.Díaz) Cuatrec. E. meridensis (Cuatrec.) Cuatrec.; VE; 2900-3200 m E. muiska (Cuatrec.) Cuatrec.; CO; 3000-3800 m E. pannosa (Standl.) Cuatrec.; VE; 3200-4150 m E. petiolata (Cuatrec.) Cuatrec.; CO; 3000-3650 m E. pleiochasia (Cuatrec.) Cuatrec.; CO; 2950-3350 m; ineludes var. pleiochasia and var. socotana Cuatrec. E. pozoensis (Cuatrec.) Cuatrec.; VE; 2700-3400 m E. purpurascens (Cuatrec.) Cuatrec.; CO, VE; 2800--3500 m E. santanderensis (A.C.Sm.) Cuatrec.; CO; 2600-4300 m E. sclerophyl!a (Cuatrec.) Cuatrec.; CO; 3200-3700 m E. trianae (Cuatrec.) Cuatrec. (syn.: Espeletia trianae Cuatrec.); CO; 3000 m Eupatorium L. (see Ageratina) Facelis Cass. F. lasiocarpa (Griseb.) Cabrera; PE; 2200-3500 m Flosca/dasia Cuatrec. F. hypsophila Cuatrec.; CO, EC; 4100-4550 m Flosmutisia Cuatrec. Re f.: Cuatrecasas, 1985b. F. paramicola Cuatrec.; CO; 3480 m Freya V.M.Badillo F. alba V.M.Badillo; VE; ca. 3200 m Galinsoga Ruiz & Pav. G. mandonii Sch.Bip.; PE; 2900--3800 m G. parviflora Cav.; CO, EC, PE; 1400-3100 m Gamochaeta Wedd. Ref.: Dillon & Sagástegui, 1986b. G. americana (Mil l.) Wedd. [syn.: Gnaphalium americanum Mill., Gn. spicatum Lam.; Gamochaeta spicata (Lam.) Cabrera]; CR, CO, VE, EC, PE; 2300-4100 m G. badi/lana (Aristeg.) A.Anderb.; VE; 3000 m G. filaginea (DC.) Cabrera (syn.: Gnaphalium filagineum DC.); EC?, PE; 3800 m
[VOL. 84
humi/is Wedd.; EC?, PE; 3600 m meridensis V.M.Badillo; VE; 3500 m oreophi/a M.O.Dillon & Sagást.; PE; 3100-4870 m paramora (S.F.Blake) A.Anderb. [syn.: Gnapha/ium paramorum S.F.Blakef Lucilia paramorum (S.F.Blake) V.M.Badillo]; CO, VE; 2900-4000 m G. pennsylvanica (Willd.) Cabrera; VE, PE; 3050-4100 m G. purpurea (L.) Cabrera (syn.: Gnaphalium columbianum Hieron.); CO, VE, EC; 3000-4300 m G. sphace/ata (Kunth) Cabrera (syn.: Gnaphalium / sphacelatum Kunth); VE, EC, PE; 3500-3600 m / G. stachydifolia (Lam.) Cabr. (syn.: Gnaphalium stachydifolia Lam.); VE; ? m G. G. G. G.
Gnaphaliothamnus Kirp. · Ref.: Nesom, 1990a, 1990b, 1994a. G. /avendulaceum G.L.Nesom (DC.) [syn.: Gnaphalium lavandulaceum DC., G. lavandulifolium (Kunth) S.F.Blake; Chionolaena /avandulifo/ia (Kunth) S.F.Blake]; CR; 3100·-3800 m Gnapha/ium L. (see also synonyms under Achyrocline, Be/loa, Chionolaena, Gamochaeta, Lucilia) G. antennarioides DC.; CO, VE, EC, PE; 3400-4300 m G. attenuatum DC.; CR, PA; 2900-3200 m G. cheiranthifolia Lam.; CO, VE, EC; 3200-4200 m G. chimborazense Hieron.; EC; 4000-4300 m G. dombeyanum DC. [syn.: Pseudognaphalium dombeyanum (DC.) A.Anderb.]; CO, VE, EC, PE; 3300-4400 m G. dysodes Spreng.; EC; 3000-3550 m G. e/egans Kunth [syn.: Pseudognaphalium elegans (Kunth) A.Anderb.; G. poeppigianum DC.]; CO, VE, EC, PE; 3000-3450 m G. gnaphalioides (Less.) P.Beauv.; CO; 3650-3900 m G. lanuginosum Kunth; CO?, PE; 3500 m G. liebmannii Sch.Bip. ex Klatt.; CR; 3000-3500 m; includes var. monticola (McVaugh) D.Nash G. linearum Cuatrec.; CO; 3000-3500 m G. meridanum Aristeg. [syn.: Pseudognaphalium meridanum (Aristeg.) A.Anderb.]; CO, VE, EC; 3500-4500 m G. polium Wedd.; PE; 3600-3800 m G. rhodarum S.F.Blake; CR, PA; 2700-3800 m G. roseum Kunth; CR; 2000-3500 m G. rosulatum S.Moore; CO; 3100-4900 m G. schlimii Briq.; CO; 2450-3050 m G. subsericeum S.F.B!ake; CR, CO, PE; 2800 m G. tenue Kunth; CO, EC; 2300-3500 m Grosvenoria R.M.King & H.Rob. G. hypargyra (B.L.Rob.) R.M.King & H.Rob.; EC; 30003800m G. rimbachii (B.L.Rob.) R.M.King & H.Rob.; EC; 35003700 m Gy1wxys Cass. [Note: Many trees of this genus are found in protected habitats with Polylepis.] Ref.: Herrera A. de Loja, 1980.
1~9]
PARAMOS
G. acostae Cuatrec.; EC; 3200-4000 m G. albivestita Cuatrec.; CO; 3450-4300 m G. apollinaris Cuatrec.; CO; 3600-3700 m G. arnicae Cuatrec. var. scandens Cuatrec.; CO; 3000-3600 m G. baccharoides (Kunth) Cass.; EC; 3900-4100 m G. bracteolatr; Cuatrec.; CO; 3600-4400 m G. buxifolia (Kunth) Cass.; CO, EC, PE; 3000-4100 m G. calyculisolvens Rieron.; EC, PE; 3100-3550 m G. caracensis Muschl.; PE; 3400-4000 m G. chingualensis R.Rob. & Cuatrec.; EC; 3400-3600 m G. colanensis M.O.Dillon & Sagást.; PE; 3000-3450 m G. columbiana (K1att) Rieron.; CO; ? m J G. costihirsuta Cuatrec.; PE; 3200-3450 m G. cuicochensis Cuatrec.; EC; 3100-3500 m G. dilloniana Sagást. & Tellez; PE; 3300-4100 m G.ferreyrae B.Rerrera; PE; 3400-3850 m 1 G. jlexopedes Cuatrec.; CO; 3000-3500 m G.jlorulenta Cuatrec.; CO; 3000-3500 m G. hallii Rieron.; EC; 3300-3900 m .G. hirsuta Wedd.; CO; 3450-4000 m G. hutchisonii R.Rob. & Cuatrec.; PE; 3400-3550 m G. hypoleucophy/la Cuatrec.; PE; 3200-3450 m G. induta Cuatrec.; CO; 3400-3550 m G. laurata Cuatrec.; CO; 3400-3500 m G. lehmannii Rieron.; CO; 3300-3600 m G. lindenii Sch.Bip.; CO; 3700 m G. meridana Cuatrec.; VE, EC; 3050-4100 m G. miniphylla Cuatrec.; EC; 3100-3950 m G. moritziana Sch.Bip.; VE; 3100-3800 m G. nítida Muschl.; PE; 3500-3850 m G. paramuna Cuatrec.; CO; 3750-4100 m • G. parvifolia Cuatrec.; CO, EC; 3000-4100 m _ G. pendula Sch.Bip. ex·Wedd.; CO, EC; 3600-4200 m .G. perbracteata Cuatrec.; CO, EC; 3400 m G. rugulosa Muschl.; PE; 3700-4100 m G. sancti-antonii Cuatrec.; CO, EC; 3200-4200 m G. sodiroi Rieron.; EC, PE; 3400-3800 m G. soukupii ~uatrec.; PE; 2800-3550 m G. subcinerá Cuatrec.; CO; 3000-4200 m G. subhirsuta Cuatrec.; CO; 3650-3800 m G. tolimensis Cuatrec.; CO, EC; 3350-4300 m G. tomentosissima Cuatrec.; PE; 3000-3500 m G. trianae Rieron.; CO; 3400-4000 m G. validifolia Cuatrec.; EC; 31 00-3'450 m G. violacea Sch.Bip. ex Wedd.; VE; 2500-3850 m G. visoensis Cuatrec.; PE; 3800 m
91
'Hieracium L. H. avilae Kunth; CO, VE, EC; 3100-4050 m H. eriadenium S1eumer; PE; 2000-4000 m H. erianthum Kunth; CO, VE; 3600-3800 m H.frigidum Wedd.; CO, VE, EC, PE; 3250-4900 m H. irasuense Benth. (syn.: H. maxonii S.F.Blake, H. standleyi S.F.Blake); CR, PA; 1500-3800 m
H. H. H. H. H. H. H. H.
lagopus D.Don; EC, PE; 3100-3250 m lehmannii Zahn; CO; 3550-3700 m mapirense Britton; PE; 2800-3500 m paletaranum S1eumer; CO; 3100-3250 m sodiroanum Zahn; EC; 2800-3400 m tal/enganum Zahn; PE; 1900-3750 m tolimense Cuatrec.; CO; 3000-4200 m venezuelanum Arv.-Touv.; VE; 3600 m
Hiliterhubera Sch.Bip. ex Wedd. Ref.: Cuatrecasas & Aristeguieta, 1956.
H. H. H. H. H. H. H. H.
adenopetala Cuatrec. & Aristeg.; VE; 3200-4200 m columbica Sch.Bip. ex Wedd.; CO, VE; 3800-4900 m ericoides Wedd.; VE; 3500-4300 m harrietae Cuatrec.; CO; 4500-4900 m imbrica/a Cuatrec. & Aristeg.; VE; 3900-4000 m lanuginosa Cuatrec. & Aristeg.; VE; 3500-4300 m laseguei Wedd.; VE; 3700-4800 m nevadensis Cuatrec.; CO; 3500-4850 m
Hymenostephium Benth. (see Viguiera) Hypochaeris L. (not "Rypochoeris") H. barbato (Sch.Bip.) Reiche; CO, EC, PE; 3750-4450 m H. elata Griseb.; EC, PE; 2600-3150 m H. eriolaena (Sch.Bip.) Reiche; PE; 3200-3850 m H. gramínea Rieron.; PE; 2400-3500 m H. hohenackeri Sch.Bip.; PE; 3500-3750 m H. meyeriana (Wa1p.) Griseb.; EC, PE; 2500-3500 m H. radica/a L.; CR, CO, EC; 1800-4000 m H. sessilijlora Kunth; CO, VE, EC, PE; 3450-4900 m; ineludes var. albiflora Rieron. and var. sessilijlora H. setosa (Wedd.) Rusby; CO, VE, EC, PE; 3200-4100 m H. sonchoides Kunth; EC; 3500-4 700 m H. taraxacoides (Walp.) Benth. & Rook.f.; CO, PE; 30004300m
Iltisia S.F.Blake l echandiensis R.M.King & R.Rob.; CR, PA; 3000-3550 m l repens S.F.Blake; CR; 3100-3500 m
Haplopappus Cass. (see Llerasia)
Jalcophila M.O.Dillon & Sagásteg.
Helianthopsis R.Rob. (see Pappobolus)
Ref.: Díllon & Sagástegui, 1986a. J. ecuadoriensis M.O.Dillon & Sagásteg.; EC; 3750-4000 m J. peruviana M.O.Dillon & Sagásteg.; PE; 3250-3800 m
Helianthus L. (see Pappolobus) Heliopsis Pers. H. decumbens S.F.B1ake; PE; 3100 m H. lanceo/ata S.F.B1ake; CO; 2800-3450 m
Jaramil/oa R.M.King & R.Rob. J. hylibates (B.L.Rob.)R.M.King & R.Rob.; CO; 2700-3750 m Jungia L.f. Ref.: Rar1ing, 1991.
Helogyne Nutt. H. calocephala Mattf.; PE; 3300-3400 m
J. coarctata Rieron. (syn.: J. lehmannii Rieron.); CO, EC; 2800-3600 m
92
.MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
J.jerruginea L.f.; CO, VE, EC; 2500-3650 m J. karstenii Sch.Bip. ex Cuatrec.; CO; 4000 m , J. panicu/ata (DC.) A.Gray subsp. stipu/ifera (Cuatrec.) Har1ing; CO; 2500-3300 m J. rugosa Less.; EC, PE; 2800-3700 m J. stuebe/ii (Hieron.) Crisci; PE; 2200-3500 m Laestadia Kunth ex Less. L. costaricensis S.F.B1ake; CR, PA; 3000-3500 m L. muscico/a Sch.Bip. ex Wedd.; CO, VE, EC, PE; 32004050m L. pinifolia Kunth ex Less.; CO; 3150-3900 m L. rupestris Benth.; CO; 3400-3800 m
[VOL. 84
L. neriifo/ius (Sch.Bip./ex Wedd.) Emst.; CO, VE; 20003600 m; inc1udes var. neriifolius and var. turmalensis Cuatree, L. occultus (S.F.Blake) Cuatrec. [syn.: L. glossophyllus (Mattf.) Cuatrec., 'L. humbertii (Cuatrec.) Cuatrec.]; CO, VE; 2700-4000 m; includes var. occultus and var. salomonii Cuatrec. & López-Fig. L. parvulus Cuatrec.; VE; 3100-3400 m L. spectabilis Cuatrec.; VE; 2900-3250 m L. tam~nus (Cuatrec.) Cuatrec. (syn.: Espeletia tamanus Cuatrec.); CO; 2900-3300 m
Llerasia Triana [Note: Mostly a forest genus, but sorne species are found . in disturbed páramo.] Lagenifera Cass. (see Myriactis) . ·L. caucana (S.F.B1ake) Cuatrec.; CO; 2500-3600 m L. hypoleuca (Turcz.) Cuatrec. var. serrata Cuatrec. (syn.: HapLagenophora Cass. (syn.: Lagenifera Cass.) (seeMyriactis) p/opapus hypoleucus Turcz.); CO, EC; 1580-3700 m Lasiocepha/us Willd. ex Sch1tdl. (syn.: Aetheolaena Cass ..) L.lindeniiTriana var. subcalva (S.F.B1ake) Cuatrec. (syn.: Happ/opapus lehmannii S.F.B1ake); CO, EC; 2700-3600 m Ref.: Cuatrecasas, 1978. L. cuencanus (Hieran.) Cuatrec.; CO, VE, EC; 2950-3600 m L. doryphyllus (Cuatrec.) Cuatrec.; CO, VE; 3850-4400 m Loricaria Wedd. Re f.: Cuatrecasas, 1954. L. gargantanus (Cuatrec.) Cuatrec.; CO, EC; 3800-4600 m L. heterophyllus (Turcz.) Cuatrec.; EC; 2900-3300 m L. antisanensis Cuatrec.; EC; 3950-4200 m L. involucratus (Kunth) Cuatrec.; CO, EC, PE; 3200-4150 m L. azuayensis Cuatrec.; EC; 3800-3950 m L. co/umbiana Cuatrec.; CO; 3500-5000 m L. lingulatus Schltdl. [syn.: Senecio lingulatus (Schltdl.) L. complanata (Sch.Bip.) Wedd. subsp. occidentalis Cuatrec.]; EC; 3450-4200 m Cuatrec.; CO, EC; 3350-4500 m; inc1udes var. occiL. /oeseneri (Hieron.) Cuatrec.; PE; 2400-3450 m dentalis Cuatrec. and var. tatamensis Cuatrec. L. longepenicillatus (Sch.Bip. ex Sandwith) Cuatrec. (syn.: L.ferruginea (Ruiz & Pav.) Wedd.; EC, PE; 3100-4800 m Senecio /ongepenicil/atus Sch.Bip. ex Sandwith, S. pittierii Cuatrec.); VE; 2900-4400 m L. ilinissae (Benth.) Cuatrec.; EC; 3500-4450 m L. lagunillensis Cuatrec.; CO; 3500-3800 m L. mojandensis (Hieron.) Cuatrec.; CO, EC; 3200-4400 m L. otophorus (Wedd.) Cuatrec. (syn.: Senecio otephorus L. leptothamna (Mattf.) Cuatrec.; PE; 3400-4000 m L. lycopodjnea Cuatrec.; PE; 4000-4100 m Wedd.); CO, EC; 3350-4400 m L. macbridei Cuatrec.; PE; 4000 m L. ovatus Sch1tdl. [syn.: Culcitium ovatum (Schltdl.) S.F.Blake, C. reflexum Kunth, C. uniflorum (Lam.) L. oel/gaardii M.O.Dillon & Sagást.; EC; 3350-3450 m Hieron.; Senecio superandinus Cuatrec.]; CO, EC; 3200L. pauciflora Cuatrec.; PE; 3350 m 4500m L. puracensis Cuatrec.; CO, EC; 3500-3600 m L. patens (Kunth) Cuatrec. (syn.: Gynoxys auriculata L. sco/opendra (Hook.) Kuntze; EC; 3050-4200 m Turcz.); CO, VE, EC, PE; 3600-4300 m L. thuyoides (Lam.) Sch.Bip.; CO, EC, PE; 3300-4500 m; L. puracensis (Cuatrec.) Cuatrec. (syn.: Culcitium pura cense inc1udes var. stuebelii (Hieron.) Cuatrec. and var. Cuatrec.); CO; 3800-4500 m thuyoides L. sodiroi (Hieran.) Cuatrec.; EC; 3200-4300 m Lourteigia R.M.King & H.Rob. Liabum Adans. (see Munnozia senecioides) L. bal/otaefo/ia (Kunth) R.M.King & H.Rob.; CO; 27003750m L. humilis (Benth.) R.M.King & H.Rob. [syn.: Eupatorium Libanothamnus Ernst humile (Benth.) Hieron.]; CO; 3150-3700 m [Note: A forest genus that reaches into páramo under disL. lanulata (B.L.Rob.) R.M.King & H.Rob.; CO; 2800-3600 m turbance or edaphic factors.] L. arboreus (Aristeg.) Cuatrec.; VE; 3100-3350 m L. microphylla (L. f.) R.M.King & H.Rob.; CO; 3150-3600 m L. banksiaefolius (Sch.Bip. & Ettingsj. ex Wedd.) Cuatrec.; L. ornatiloba (B.L.Rob.) R.M.King & H.Rob. (syn.: Eupa. torium ornatilobum B.L.Rob.); CO; 2800-3800 m VE; 2700-3300 m; includes var. banksiaefolius and var. L. scandens V .M.Badillo; VE; 3100 m granatensis Cuatrec. L. divisoriensis Cuatrec.; CO, VE; 3200-3650 m L. stoechadifo/ia (L. f.) R.M.King & H.Rob. (syn.: Eupatorium stoechadifolium L.f.); CO, VE; 2900-4100 m L. griffinii (Ruiz-Terán & López-Fig.) Cuatrec.; VE; 26003100m Lucilia Cass. L. liscanoanus (Cuatrec.) Cuatrec.; VE; 2900-:-3550 m L. kunthiana (DC.) Zardini [syn.: Gnaphalium kunthianum L. lucidus (Aristeg.) Cuatrec.; VE; 3300-3500 m
1999]
PARAMOS
(DC.) Kuntze, L. lehniannii Hieran., L. pusilla Hieran., . L. venezuelensis Steyerm.]; CO, VE, EC, PE; 36504500 m L. saxatilis V.M.Badillo; VE; 3000-3100 m Luciliopsis Wedd. (see Cuatrecasasiella) Matricaria L. [Note: This genus is cu1tivated as an herbal and rare1y escapes into páramo.] M chamomilla L. [syn.: M recutita L.; Chamomilla recutita (L.) Rauschert]; CO; 3500-3600 m Mikania Willd. [Note: Most1y a forest genus, but reaching páramo in disturbed situations.] M aristei B.L.Rob.; CO; 2300-3100 m M caldasana B.L.Rob.; CO; 3000-3500 m M harlingii R.M.King & H.Rob.; EC; 2700-3000 m M iltisii R.M.King & H.Rob.; CR; 1900-3300 m M miconoides B.L.Rob.; VE; 2700-3300 m M. stuebelii Hieran.; CO, EC; 2700-3700 m M violascens (B.L.Rob.) R.M.King & H.Rob.; CO; 29003250 m Mniodes (A.Gray) A.Gray ex Benth. & Hook.f. M pulvinata Cuatrec.; PE; 3900-4900 m Monactis Kunth M macbridei H.Rob.; PE; 2800-3600 m Monticalia C.Jefftey (see species in synonymy of Pentacalia) Munnozia Ruiz & Pav. M acostae (I Chung) H.Rob. & Brettell; EC; 3200-4000 m Mjussieui (Cass.) H.Rob. & Brettell (syn.: Liabum nonoense Hieran., L. tabanense Cuatrec.); CO, EC; 25003800m M lyrata (A.Gray) H.Rob. & Brettell; PE; 2800-3800 m M. nívea (Hieran.) H.Rob. & Brettell; CO, EC, PE; 26003500m M peruensis (Cuatrec.) H.Rob. & Brettell; PE; 1800-3500 m M. sagittata (Sch.Bip.) H.Rob. & Brettell (syn.: Liabum sagittatum Sch.Bip.); CO; 2850 m M. senecionidis Benth. (syn.: Liabum megacephalum Sch.Bip.); CO, V~ 3200-3550 m Mutisia L. f. Ref.: Cabrera, 1965. M afata Hieran.; EC; 2900-3900 m M. andersonii Sodira ex Hieran.; EC; 3000-3500 m M clematis L. f.; CO, EC; 2600-3900 m M intermedia Hieran.; EC; 2000-3400 m [Note: Incorrect1y cited by Harling (1991) as M. grandiflora Humb. & Bonpl. (Ulloa U. & Jorgensen, 1996).] M. lehmannii Hieran.; EC; 3500-3700 m M microcephala Sodira ex Cabrera; EC; 3300-3800 m M. microphylla Willd. ex DC.; EC; 2850-4180 m
93
M. ochroleucá Cuatrec.; CO; 3100-3500 m M. sodiroi Hieran.; CO, EC; 2750-3650 m M. viridis Cuatrec.; CO; 3450-3500 m Myriactis Less. [syn.: Lagenophora Cass. (= Lagenifera Cass.)] M. andina (V.M.Badillo) M.C.Vélez (syn.: Lagenophora andina V.M.Badillo); CR, VE; 3600-4400 m M. minuscula (Cuatrec.) Cuatrec. (syn.: Lagenophora minuscu.la Cuatrec.); PA; 3150-3335 m M. panamensis (S.F.B1ake) Cuatrec. (syn.: Lagenophora panamensis S.F.B1ake); CR, PA; 2800-3400 m M. sakirana (Cuatrec.) Cuatrec. (syn.: Lagenifera sakirana Cuatrec.); CR; 3000-3500 m M westonii (Cuatrec.) Cuatrec. (syn.: Lagenifera westonii Cuatrec.); CR; 3200-3500 m Noticastrum DC. N. marginatum (Kunth) Cuatrec. (syn.: Aster marginatus Kunth); CO, VE, PE; 3550-4200 m Novenia S.E.Freire N. acaulis (Wedd. ex Benth.) S.E.Freire & Hellweg; PE; 3100-4600 m Oritrophium (Kunth) Cuatrec. (syn.: Ce/misia Cass.) Ref.: Cuatrecasas, 1961, 1969, 1997. O. blepharophyllum (S.F.Blake) Cuatrec. (syn.: Erigeron blepharophyllum S.F.Blake); VE; 3000-3600 m O. cocuyense (Cuatrec.) Cuatrec. (syn.: Erigeron cocuyensis Cuatrec.); CO, EC; 3350-4400 m O. crocifolium (Kunth) Cuatrec. [syn.: Ce/misia crocifolia (Kunth) Sch.Bip.]; EC; 3400-4300 m O.figueirasii Cuatrec.; VE; 3300 m O. granatum Cuatrec.; VE; 3400-3750 m O. hieracioides (Wedd.) Cuatrec. [syn.: O. ferrugineum (Wedd.) Cuatrec.]; PE; 2700-3700 m O. hirtopilosum (Hieran.) Cuatrec.; PE; 2900-3650 m O. limnophilum (Sch.Bip.) Cuatrec.; CO, VE, EC, PE; 34004 700 m; in eludes subsp. limnophilum, subsp. mutisianum (Cuatrec.) Cuatrec., subsp. nevadanum Cuatrec., and subsp. punae Cuatrec. O. mucidum Cuatrec.; CO; 3000-3200 m O. nevadense (Wedd.) Cuatrec. (syn.: Erigeron nevadensis Wedd.); VE; 3500-4100 m O. oellgaardii Cuatrec.; EC; 3550-4000 m O. peruvianum (Lam.) Cuatrec. [syn.: Ce/misia peruviana (Lam.); Erigeron peruvianus (Lam.) Cuatrec.; Aster pellitus Kunth]; CO, VE, EC, PE; 3300-4600 m; includes subsp. lineatum (Cuatrec.) Cuatrec., subsp. perijaense Cuatrec., subsp. peruvianum fo. peruvianum Cuatrec. and fo. intermedium Cuatrec. O. repens (Kunth) Cuatrec.; EC, PE; 3100-3400 m O. tergoalbum (Cuatrec.) Cuatrec. (syn.: Erigeron tergoalbus Cuatrec.); EC; 3050-3400 m O. venezuelense (Steyerm.) Cuatrec. (syn.: Erigeron venezuelense Steyerm.); VE; 3250-3700 m
94
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
O:zy/obus (Moc. ex DC.) A.Gray Ref.: Tumer & Kerr, 1985. O. glandu/iferus (Sch.Bip. ex Benth. & Hook.f.) A.Gray (syn.: Ageratum sordtdum S.F.B1ake); CO, VE; 30504070m Pappobolus S.F.B1ake (syn.: Helianthopsts H.Rob.) Ref.: Panero, 1989. P. decumbens Panero; PE; 2850-3350 m P. dtsco/or (S.F.B1ake) Panero; PE; 2900-350Cl m P.je/sktt (Hieran.) Panero; PE; 3050-3500 m P. lodtcatus (Cuatrec.) Panero (syn.: Helianthus lodtcatus Cuatrec.); PE; 3000-3700 m P. mtcrophyllus (Kunth) Panero var. dillontt Panero; PE; 2600-3400 m P. smtthti (Ferreyra) Panero; PE; 3300-3500 m P. stuebelli (Hieran.) Panero; PE; 3000-3600 m P. subniveus (S.F.Blake) Panero (syn.: Heltanthus subntveus S.F.B1ake); PE; 2800-3700 m Paramiflos Cuatrec. Re f.: Cuatrecasas, 199 5. P. glandulosus (Cuatrec.) Cuatrec. [syn.: Espe/etiopsis glandu/osus (Cuatrec.) Cuatrec.]; CO; 3000-3500 m Paranephelius Poepp. P.ferreyrti H.Rob.; PE; 3100-3 750 m P.je/skii (Hieran.) H.Rob. & Brettell; PE; 2200-3750 m P. ovatus Wedd.; PE; 3300-4000 m P. unijlorus Poepp. & Endl.; PE; 3200-3800 m P. wurdackii H.Rob.; PE; 3500-3750 m Pentacalia Cass. (syn.: Monticalia C.Jeffrey) Ref.: Cuatrecasas, 1981; Robinson & Cuatrecasas, 1993. P. abietina (Willd. ex w·edd.) Cuatrec. [syn.: Monticalia abtetina (Willd. ex Wedd.) C.Jeffrey; Senecio aciculatus Cuatrec.]; CO, VE; 3100-4200 m; includes var. abietina and var. acicula/a (Cuatrec.) Cuatrec. P. aclydtphylla (Cuatrec.) Cuatrec. (syn.: Senecto ac/ydiphyllus Cuatrec.); PE; 3400-3750 m P. albi-panquei S.Díaz & Cuatrec.; CO; 3140-3400 m P. albotecta (Cuatrec.) Cuatrec. (syn.: Senecio albotectus Cuatrec.); CO; 3000-3600 m P. americana (L.f.) Cuatrec.; CO; 2600-4100 m P. amplexicaulis (Kunth) Cuatrec. (syn.: Senecio amplexicau/is Kunth; Dendrophorbium amp/exicaulis (Kunth) B.Nord.); EC; 2400-3100 m P. andico/a (Turcz.) Cuatrec. [syn.: Montica/ia andico/a (Turcz.) C.Jeffrey; Senecio andicolus Turcz.]; CR, PA, CO, VE, EC, PE; 2900-4400 m P. antioqutensis (Cuatrec.) Cuatrec.; CO; 1500-3100 m P. apiculata (Sch.Bip. ex Wedd.) Cuatrec. [syn.: Monticalia apiculata (Sch.Bip. ex Wedd.) C.Jeffrey; Senecio apiculatus Sch.Bip. ex Wedd.]; VE; 3300-3500 m P. arborea (Kunth) H.Rob. & Cuatrec.; CO, EC; 27003400 m P. arbutifolia (Kunth) Cuatrec. (syn.: Senecio arbutifolius Kunth, S. pichtnchensis Greenm.); CO, EC; 3600-4500 m P. batallonensis (Cuatrec.) Cuatrec. [syn.: Monticalia
, [VOL. 84
batallonensis (Cuatrec.) C.Jeffrey; ;Jenecio batallonensis Cuatrec.]; VE; 3400 m ~ P. befariotdes (Cuatrec.)Cuatrec. [syn.: Senecio befariotd s Cuatrec.; Monticalia befariotdes (Cuatrec.) B:Nord. ; EC; 3400-4500 m P. breviligu/ata (Hieran.) Cuatrec.; CO; 3000-3390 m P. cacaosensis (Cuatrec.) Cuatrec. (syn.: Senecio cacaosensis Cuatrec.); CO; 3100-3450 m P. cachacoensis (Cuatrec.) Cuatrec. [syn.: Monticalia cachacoensis (Cuatrec.) C.Jeffrey; Senecio cachacoensis Cuatrec.]; CO, VE; 3200-3700 m P. campii (Cuatrec.) Cuatrec.; EC; 3100-3550 m P. carrikeri (Cuatrec.) Cuatrec. (syn.: Senecio carrikeri Cuatrec.); CO; 3200-3700 m P. chaquiroensis (Greenm.) Cuatrec. (syn.: Senecio chaquiroensis Greenm.); CO; 3100-3600 m P. cleefit (Cuatrec.) Cuatrec. (syn.: Senecto cleefii Cuatrec.); CO; 4200-4400 m P. colombiana (Cuatrec.) Cuatrec. (syn.: Senecio co/ombtanus Cuatrec.); CO, VE; 3200-3900 m P. corazonensts (Hieran.) Cuatrec.; EC; 2800-3550 m P. corymbosa (Benth.) Cuatrec. (syn.: Senecio corymbosus ~enth.; S. microchaete Wedd., nom. illeg.); CO; 28503350m P. dtamantensts (Cuatrec.) Cuatrec,; CO; 3000-3600 m P. elatoides (Wedd.) Cuatrec.; CO;? m P. el/tpticifolta (Hieran.) Cuatrec.; CO; 1750-3200 m P. empetroides (Cuatrec.) Cuatrec. (syn.: Senecio empetrotdes Cuatrec.); CO, EC; 3100-3500 m P. firmipes (Greenm.) Cuatrec. (syn.: Senecio firmipes Greenm.); CR, PA; 3200-3500 m P.j/occultdens (Sch.Bip. ex Wedd.) Cuatrec. [syn.: Monticaliaj/occulidens (Sch.Bip. ex Wedd.) C.Jeffrey; Senecio jlocculidens Sch.Bip. ex Wedd.]; VE; 2800-3300 m P. flosfragrans (Cuatrec.) Cuatrec. (syn.: Senecio j/osfragrans Cuatrec.); CO; 3500-4425 m P. gelida (Wedd.) Cuatrec. (syn.: Senecio geltdus Wedd.); CO; 4200-4700 m P. greenmaniana (Hieran.) Cuatrec. [syn.: Monticalia greenmaniana (Hieran.) C.Jeffrey; Senecto greenmantanus Hieran.]; VE; 3000-4100 m P. guadalupe (Cuatrec.) Cuatrec. (syn.: Senecto guadalupe Cuatrec.); CO; 3350-4000 m P. guicanensis (Cuatrec.) Cuatrec. (syn.: Senecio gutcanensts Cuatrec.); CO; 4000-4400 m P. harrietae (Cuatrec.) Cuatrec.; CO; 3350 m P. tmbricatifolia (Sch.Bip. ex Wedd.) Cuatrec. [syn.: Montica/ia imbrtcatzfolia (Sch.Bip. ex Wedd.) C.Jeffrey]; VE; 4265-3900 m P. karstenti (Hieran.) Cuatrec. (syn.: Senecio karstenii Hieran.); CO; ? m P. kleinioides (Kunth) Cuatrec.; CO; 1900-3300 m P. /anceolifolia (Cuatrec.) Cuatrec.; EC; 3400 m P. ledifolta (Kunth) Cuatrec. [syn.: Monttcalia ledzfolia (Kunth) C.Jeffrey; Senecto ledifoltus (Kunth) DC.]; CO, VE; 3200-3850 m; inc1udes subsp. ledzfolia, subsp. lehmannii (Rieron.) Cuatrec., and subsp. schlimti {Wedd.) Cuatrec.
;.·
1999]
PARAMOS
P. leioclada (Cuatrec.) Cuatrec. (syn.: Senecio leioclados Cuatrec.); CO; 3200-3400 m P. lindenii (Sch.Bip. ex Wedd.) Cuatrec.; CO; 2900--3800 m P. magnicalycu/ata (V.M.Badillo) Cuatrec. [syn.: Monticalia magnicalyculata (V.M.Badillo) C.Jeffrey; Senecio magnicalyculatus V.M.Badillo]; VE; 3300-3650 m P. masonhalei (Ruiz-Terán & López-Fig.) Cuatrec. [syn.: Monticalia masonhalei (Ruiz-Terán & López-Fig.) C.Jeffrey]; VE;? m P. microdon (Wedd.) Cuatrec. [syn.: Senecio microdon Wedd.; Monticalia microdon (Wedd.) B.Nord.]; EC; 4100--4800 m P. micropachyphylla (Cuatrec.) Cuatrec. [syn.: Monticalia micropachyphylla (Cuatrec.) C.Jeffrey]; VE; 3200--3930 m . P. mutisii (Cuatrec.) Cuatrec. (syn.: Senecio mutisii Cuatrec.); CO, EC; 3300-4200 m P. myrsinites (Turcz.) Cuatrec. (syn.: Senecio ecuadoriensis Rieron.); EC; 3600-3900 m P. nigrosÍagnosa (Cuatrec.) Cuatrec. [syn.: Monticalia nigrostagnosa (Cuatrec.) C.Jeffrey; Senecio nigrostagnosus Cuatrec.]; VE; 3600-4300 m P. nítida (Kunth) Cuatrec. (syn.: Senecio nitidus Kunth); CO, EC?; 3200_:_3600 m P. novola¡;¡,ata (Cuatrec.) Cuatrec. (syn.: Senecio novolanatus Cuatrec.); CO; 3050-3450 m P. onae (Cuatrec.) Cuatrec.; EC; 2950-3050 m · P. pachypus (Greenm.) Cuatrec. [syn.: Monticalia pachypus (Greenm.) C.Jeffrey; Senecio pachypus Greenm.]; VE; 3800-4130 m P. perijaensis (Cuatrec.) Cuatrec. [syn.: Montica/ia perijaensis (Cuatrec.) C.Jeffrey; Senecio perijaensis Cuatrec.]; CO, VE; 3000-3100 m P. peruviana (Pers.) Cuatrec. (syn.: Senecio peruvianus Pers.); CO, EC, PE; 3300-4400 m P. phanerandra R.Rob. & Cuatrec.; PA; 3100-3300 m P. polymera (K1att) Cuatrec. [syn.: Monticalia polymera (K1att) C.Jeffrey]; CO, VE; 2400-3600 m P. popayanensis (Rieron.) Cuatrec.; CO, EC; 2800-3500 m P. pulchella (Kunth) Cuatrec. [syn.: Senecio pulchellus (Kunth) DC., S. pungens (Kunth) DC., S. guantivanus Cuatrec.]; CO; 3100-3700 m; inc1udes subsp. guantivanus (Cuatrec.) Cuatrec. with var. pungens (Kunth) Cuatrec. and subsp. pulchella P. quirorana (Cuatrec.) Cuatrec. [syn.: Monticalia quirorana (Cuatrec.) C.Jeffrey; Senecio quiroranus Cuatrec.]; VE; 3200-3650 m P. ramentosa (Cuatrec.) Cuatrec. [syn.: Monticalia ramentosa (Cuatrec.) C.Jeffrey; Senecio ramentosus Cuatrec.]; CO, VE; 3200--3900 m P. reissiana (Rieron.) Cuatrec. (syn.: Senecio reissianus Rieron.); CO; 3400-3900 m P. rex (Sandwith) Cuatrec. [syn.: Culcitium rex Sandwith; Monticalia rex (Sandwith) C.Jeffrey]; VE; 3800-4100 m P. ricoensis (Cuatrec.) Cuatrec.; CO; 3600 m P. rigidifo/ia (V.M.Badillo) Cuatrec. [syn.: Monticalia rigidifolia (V.M.Badillo) C.Jeffrey]; VE; 2900-3400 m P. rosmarinifolia (Benth.) Cuatrec.; EC; 3400-3800 m P. ruiteranii (Cuatrec.) Cuatrec. [syn.: Monticalia ruiteranii (Cuatrec.) C.Jeffrey; Senecio ruiteranii Cuatrec.]; VE; 3100-3400 m
95
P. scaphiformis (Oreenm.) Cuatrec.; CO; 3600-3730 m P. schultzei Cuatrec. subsp. sevillana Cuatrec.; CO; 32253450 m P. sc/erosa (Cuatrec.) Cuatrec. [syn.: Monticalia sc/erosa (Cuatrec.) C.Jeffrey; Senecio sclerosus Cuatrec.]; VE; 4000-4400 m P. stuebelii (Rieron.) Cuatrec. (syn.: Senecio stuebe/ii Rieron.); CO, EC; 3100-4150 m P. subarachnoidea (Wedd.) Cuatrec.; CO; 2400-4300 m P. summapacis (Cuatrec.) Cuatrec. (syn.: Senecio summapacis Cuatrec.); CO; 4000-4200 m P. sylvico/a (Greenm.) Cuatrec. (syn.: Senecio sylvico/us Greenm.); CO; 3700-4200 m P. taironae S.Díaz & Cuatrec.; CO; 3545-4270 m P. teretifolia (Kunth) Cuatrec. [syn.: Senecio teretifolius (Kunth) DC.; Montica/ia teretifolia (Kunth) B.Nord.]; EC, PE; 2800-3590 m P. theifolia (Benth.) Cuatrec.; CO, VE, EC; 2500-3500 m P. tolimensis (Sch.Bip. ex Wedd.) Cuatrec.; CO; 30003700 m P. trichopus (Benth.) Cuatrec. (syn.: Senecio trichopus Benth.); CO; 3400-3600 m P. tunamensis (Cuatrec.) Cuatrec. [syn.: Monticalia tunamensis (Cuatrec.) C.Jeffrey; Senecio tunamensis Cuatrec.]; VE; 3250-3600 m P. vaccinioides (Kunth) Cuatrec. [syn.: Monticalia vaccinioides (Kunth) C.Jeffrey; Senecio vaccinioides Kunth var. microdentatus Cuatrec.]; CO, VE, EC; 2900-4300 m; includes var. microdentatus Cuatrec. and var. vaccinioides P. venezuelensis (Sandwith) Cuatrec. [syn.: Monticalia venezue/ensis (Sandwith) C.Jeffrey; Senecio venezue/ensis Sandwith, S. mucuyanus Cuatrec.]; VE; 3500-4100 m P. vernicifolia (Cuatrec.) Cuatrec.; PE; 3100 m P. vernicosa (Sch.Bip. ex Wedd.) Cuatrec. (syn.: Senecio vernicosus Sch.Bip. ex Wedd.); CO, EC; 3800-4150 m P. verticillata (K1att) Cuatrec.; EC, PE; 2900-3500 m P. vicel/iptica (Cuatrec.) Cuatrec.; VE; 3300-4100 m P. viridi-alba (Cuatrec.) Cuatrec. (syn.: Senecio viridi-albus Cuatrec.); CO; 3500-3800 m P. weinmann(fo/ia (Cuatrec.) Cuatrec. (syn.: Senecio weinmannifolius Cuatrec. and fo. quindiensis Cuatrec.); CO; 3250-3500 m P. wurdackii (Cuatrec.) Cuatrec. (syn.: Senecio wurdackii Cuatrec.); PE; 3200-3500 m Perezia Lag. Ref.: Vuilleumier, 1969. P.fosbergii Tovar; PE; 3500 m [Note: Found in limestone sink ho1es.] P. multiflora (Rumb. & Bonpl.) Less.; CO, EC, PE; 32004300m P. pungens (Rumb. & Bonpl.) Less.; EC, PE; 3200-4800 m Philoglossa DC. P. mimuloides (Rieron.) R.Rob. & Cuatrec.; PE; 23003200 m [Note: This species is sometimes cultivated and escapes.]
96
MEMOIRS OF THE NEW YORK BOTANICAL OARDEN
Pfagiocheilus Am. ex DC. P.frigidus Poepp.; EC, PE; 3350--4050 m P. peduncularis (Kunth) Wedd.; EC; 3700--4300 m P. solivaeformis DC. subsp. multiflorus Cuatrec.; CO, VE, EC, PE; 3400--4300 m Podocoma Cass. (see Blakiella) Pseudognaphalium Kirp. (see Gnaphalium)
[VOL. 84
S. acama (S.F.B1ake) Longpre; CO; 2800-3700 m S. densa Longpre; CR; 3350-3600 m S. sarmentosa Less. var. papposa (S.F.B1ake) Longpre (syn.: S. triangularis var.papposa S.F.B1ake); CR; 28003300m trianae (Hieron.) Longpre (syn.: Ca/ea longipes . S.F.B1ake); CO; 3500--4100 m
s:
Scrobicaria Cass. S. aquifolia (Cuatrec.) B.Nord. (syn.: Gynoxys aquifolia
Psi/a Phi l. (see Baccharis)
Cuatrec.); VE; 2500-3200 m S. ilicifo/ia (L.f.) B.l~ord.; CO; 3550-3900 m
Raouliopsis S.F.B1ake R. pachymorpha (Wedd.) S.F.B1ake (syn.: 0/igandra pachymorpha Wedd.); VE;? m R. seifrizii S.F.B1ake; CO; 4500--4900 m Ruilopezia Cuatrec. [Note: In this genus, species followed by "(1]" are naturally found in forest, but disturbance allows them to enter páramo, according to Cuatrecasas (unpubl.).] Re f.: Cuatrecasas, 198 7. R. atropurpurea (A.C.Sm.) Cuatrec. (syn.: Espeletia afropurpurea A.C.Sm.); VE; 2600-3700 m [1] R. bracteosa (Stand!.) Cuatrec. [syn.: Espeletia bracteosa Stand!., E.frailejonota Aristeg.; R.ji·ailejonota (Aristeg.) Cuatrec.]; VE; 2600-3500 m (1] R. bromelioides (Cuatrec.) Cuatrec.; VE; 2600-2900 m [1] R. cardonae (Cuatrec.) Cuatrec. (syn.: Espeletia cardonae Cuatrec.); VE; 3000-3300 m [1] R. coloradarum (Cuatrec.) Cuatrec.; VE; 2700-2950 m [1] R. cuatrecasasii (Ruiz-Terán & López-Fig.) Cuatreé.; VE; 2500-3000 m R. emmanuelis Cuatrec.; VE; 2900-3000 m R.figueirasii (Cuatrec.) Cuatrec.; VE; 2600-3200 m [1] R. floccosa (Stand!.) Cuatrec. (syn.: Espeletia floccosa Stand!.); VE; 3000-3800 m R. grisea (Stand!.) Cuatrec. (syn.: Espeletia grisea Stand!.); VE; 3000--4000 m R. hanburiana (Cuatrec.) Cuatrec.; VE; 3100-3500 m [1] R.jabonensis (Cuatrec.) Cuatrec.; VE; 2500-3400 m R. jahnii (Stand!.) Cuatrec.; VE; 3000,--3400 m R.josephensis (Cuatrec.) Cuatrec.; VE; 2800-3300 m [1] R. leucactina (Cuatrec.) Cuatrec.; VE; 3000-3300 m R. lindenii (Sch,Bip. ex Wedd.) Cuatrec. (syn.: Espeletia lindenii Sch.Bip. ex Wedd.); VE; 2400-3400 m (1] R. lopez-palacii (Ruiz-Terán & López-Fig.) Cuatrec.; VE; 2600-3100m[1] R. marcescens (S.F.B1ake) Cuatrec. (syn.: Espeletia marcescens S.F.B1ake); VE; 2800-3200 m (1] R. margarita (Cuatrec.) Cuatrec.; VE; 3300-3400 m R. paltonioides (Stand!.) Cuatrec.; VE; 2600-3200 m [1] R. ruizii (Cuatrec.) Cuatrec.; VE; 2750-2800 m [1] R. usubillagae Cuatrec.; VE; 3000 m R. vergarae Cuatrec. & López-Fig.; VE; 2000-3200 m R. viridis (Aristeg.) Cuatrec.; VE; 2200-3600 m [1] Sabazia Cass. Ref.: Longpre, 1970.
Se/loa Kunth Ref.: Longpre, 1970; Robinson, 1997; Tumer, 1'1980. S. breviligulata Longpre [syn.: Aphanactis breviligulata · (Longpre) B.L.Turner]; CR; 3450-3800 m S. ligulata (Cuatrec.) H.Rob. (syn.: Aphanactis ligulata Cuatrec.); CO; 3600-4500 m Senecio L. (see a1so synonyms under Lasiocephalus and Pentacalia) Ref.: Vision & Dillon, 1996. S. acarinus Cabrera; PE; 2900-3600 m S. adglacialis Cuatrec.; CO; 3400-4500 m S. alberti-smithii Cuatrec.; ca·; 3950-4150 m S. almorzaderonis Cuatrec.; CO; 3100-3900 m; includes var. almorzaderonis and var. boyacensis Cuatrec. S. aridus Greenm.; CO; 3800-4300 m S. aristeguietae Cuatrec.; VE; 3200--4400 m S. ayapatensis Sch.Bip.; PE; 3350-3800 m S. canescens (Humb. & Bonpl.) Cuatrec. (syn.: Culcitium canescens Humb. & Bonpl.); CO, EC, PE; 3550-4700 m; includes var. canescens, var. macrocephalus Cuatrec., var. ecuadoriensis Cuatrec., var. elongatus Cuatrec., var. boyacensis Cuatrec., and var. magnificus Cuatrec. S. chionogeton Wedd.; CO, EC; 3200--4400 m S. chiquianensis Cabrera; PE; 2500-3800 m S. cineraria DC. (syn.: S. ginesii Cuatrec.); VE; 3500--4000 m S. coccineus K1att; CO; 3550-4000 m S. cocuyanus (Cuatrec.) Cuatrec. (syn.: Culcitium cocuyanus Cuatrec.); CO; 3800-4350 m S. collinus DC.; PE; 3300--4000 m S. comosus Sch.Bip.; PE; 4000 m S. costaricensis R.M.King; CR; 2500-3500 m S. coymolachensis Cabrera; PE; 3500-3900 m S. culcitioides Sch.Bip.; EC, PE; 3900--4600 m S. dolichodoryius Cuatrec.; PE; 3100-3400 m S.ferrugineus (K1att) Cuatrec.; EC; 3600-4500 m S. jlaccidifolius Wedd.; PE; 3200-4000 m S.folidentatus Cuatrec.; CO; 3600-3850 m S.formosissimus Cuatrec.; CO; 3300-3650 m S. formosoides Cuatrec.; CO, VE, EC; 3200--4425 m S.formósus Kunth (syn.: S. tabacon Turcz.); CO, VE, EC; 3000-4350 m S.funckii Sch.Bip.; CO, VE; 3000-4100 m S. garcibarrigae Cuatrec.; CO; 3500-4000 m S. guascensis Cuatrec.; CO; 3300-3450 m
1999]
PARAMOS
97
S. hallii Rieron.; EC; 4200-4900 m [Note: This name is a Son chus L. nom. illeg. non Britton and seeming1y has not been re[Note: A genus of European weeds now natura1ized in p1aced (P. Jorgensen, pers~ comm.).] the N eotropics.] . S. hansweberi Cuatrec.; CR; 3400-3850 m S. asper (L.) Rill; VE, PE; 3100-4100 m S. hastatifolius Cabrera; PE; 3600-3950 m S. oleraceus L.; CR, CO, VE, EC, PE; 3000-3800 m S. hohenackeri Sch.Bip.; PE; 3750 m S. hypsobates Wedd.; CO, EC; 3300-4400 m; inc1udes var. Stevia Cav. hypsobates and var. parvulus Cuatrec. S. cajabambensis Rieron.; PE; 2250-3750 m S. isabelis S.Díaz; CO; 4400-4800 m S. elatior Kunth; CO, VE, EC; 3200-3600 m S. ishcaivilcanus Cuatrec.; PE; 3800-3900 m S. incognita Grashoff; CO, VE; 3100-3400 m S. kingbishopii Cuatrec.; PE; 3300-4000 m S. lucida Lag.; CR, PA, CO, VE; 3050-4900 m S. kuhbieri Cuatrec.; CR; 3400-3800 m S. macbridei B.L.Rob.; PE; 2200-3500 m S. laricifolius Kunth; PE; 2400-3750 m S. mandonii Sch.Bip.; PE; 3500 m S. latijlorus Wedd. v·ar.lacunosus Cuatrec.; CO, EC; 3200S. ovala Willd. (syn.: S. rhombifolia Kunth); CO, VE, EC; 4800 m 2200-3600 m S. leucanthemoides Cuatrec.; CO; 2800-4400 m S.pallida (Sch.Bip.) Rieron.; CO, VE; 3200-3700 m S. lopez-mirandae Cabrera; PE; 3200 m S. triflora DC.; VE; 1100-4100 m S. minesinus Cuatrec.; PE; 3150-4500 m S. weberbaueri B.L.Rob.; PE; 3250-3800 m S. miniauritis Sagást. & M.O.Dillon; PE; 2650-3100 n\ S. westonii R.M.King & R.Rob.; CR; 3350 m s.· mitonis Cuatrec.; PE; 3600-3800 m S. multivenius Benth.; CR; 1000-3800 m Stuckertiella Beauverd S. nivalis (Kunth) Cuatrec. (syn.: Culcitium nivale Kunth); S. capitata (Wedd.) Beauverd; EC, PE; 3100-3500 m EC; 3600-4600 m S. niveoaureus Cuatrec.; CO; 3200-4300 m "'-. S. nubigenus Kunth var. pimpinelifolia (see Dorobaea) Tagetes L. '-.___S. oerstedianus Benth.; CR, PA; 2000-3800 m [Note: Many species ofthis genus are cu1tivated and/or S. pascuiandinus Cuatrec.; CO; 4000-4300 m have now escaped into páramo.] S. pleniauritus Cuatrec.; PE; 3200-3450 m T. elliptica Smith; PE; 2200-3400 m [Note: Cu1tivated.] S. pseudoformosus Cuatrec.; CO; 3550-3900 m T.filifolia Lag. (syn.: T. pusilla Kunth); VE, EC, PE; 3200S. punasessilis Cuatrec.; PE; 3600-3900 m 3500m S. rhizocephalus Turcz. [S. repens DC. auct., nom. illeg. non T. ternijlora Kunth; PE; 3200-3450 m [Note: Cultivated.] Stokes (1812)]; CO, EC, PE; 3550-4800 m S. rhizomatus Rusby; PE; 4000 m Talamancalia H.Rob. & Cuatrec. S. romeroi Cuatrec.; CO; 3500-3550 in T. westonii R.Rob. & Cuatrec. [syn.: Pseudogynoxys S. rubrilacunae Cuatrec.; CO; 3600-4200 m westonii (R.Rob. & Cuatrec.) B.L.Tumer]; CR; 3100 m S. santanderensis Cuatrec.; CO; 3_600-4100 m S. scrobicarioides DC.; PE; 4000 m Taraxacum F.R.Wigg. S. silphioides Rieron.; CO; 2500-3900 m [Note: A genus ofintroduced roadside weeds.] S. subruncinnatus Greenm.; CO, VE; 3350-3700 m T. craspedotoides Richards; VE; 3400-4900 m S. summus Cuatrec. (syn.: Culcitium longifolium Turcz.); T. officinale G.Weber; CO, VE, EC, PE; 1700-4200 m CO, EC; 3800-4400 m; includes var.longifolius (Turcz.) T. vulgare (Lam.) Schrank; CO, EC; 2200-4400 m [Note: Cuatrec. and var. summus Perhaps this species is equa1 to T. officinale.] S. supremus Cuatrec.; CO; 4300-4600 m S. tephrosioides Turcz.; CO, EC, PE; 3250-4200 m S. tergolanatus Cuatrec.; CO; 3500-3800 m Trichocline Cass. S. tingoensis Cabrera & Zardini; PE; 3200-3800 m Ref.: Ransen, 1990; Zardini, 1975. S. usgoroensis Cuatrec.; PE; 2250-3750 m T. hieracioides (Kunth) Ferreyra [syn.: Gerbera hieracioides S. vulgaris L.; VE, EC; 2200-3500 m (Kunth) Zardini; T. beckeri B.L.Rob., T. peruviana S. yauyensis Cabrera; PE; 3200 m Rieron.]; EC, PE; 2600-3500 m Sigesbeckia L. S. bogotensis D.L.Schu1z; CO, EC; 2500-3300 m 'S.jorullensis Kunth (syn.: S. cordifolia Kunth); CR, CO, VE, EC, PE; 3000-3800 m S. orientalis L.; CO, VE; 3000-:-3500 m
Tridax L. T. angustifolius Spruce ex Benth. & Rook.f. (syn.: T. sodiroi Rieron., T. stuebelii Rieron.); EC, PE; 2700-3500 m T. tambensis Rieron.; PE; 3200 m
Smal/anthus Mack. S. pyramidalis (Triana) R.Rob. (syn.: Polymnia pyramidalis Triana, P. eurylepis S.F.B1ake); EC; 3650-4600 m [Note: Often cu1tivated near bornes.]
Vasquezia Phi!. V. anemonifolia (Kunth) S.F.B1ake (syn.: Villanova anemonifolia Less.); CO, VE; 1800-3500 m V. titicacensis (Meyen & Wa1p.) S.F.Biake; CO, EC, PE; ? m
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MEMOIRS OF THE NEW YORK BOTANICAL GARDEN ·
Viguiera Kunth [Note: Doubtfully a páramo genus.) V. goebellii (Klatt) H.Rob. [syn.: Hymenostephium goebel/ii (Klatt) H.Rob.); VE; 2600-3500 m V. sodiroi (Hieron.) S.F,Blake; EC; 2800-3950 m Vil/anova Lag. (see Vasquezia) Werneria Kunth (see also synonyms under Xenophyl/um) Ref.: Fu~k, 1997a, 1997b. W. caespitosa Wedd.; PE; 3500-3900 m W. nubigena Kunth (sy-n.: W. distichia Kunth, W. stuebelii Hieron.); CR, CO, EC, PE; 3500-4400 m W. pumila Kunth; CO, EC; 3700--4800 m W. pygmaea Gillies ex Hook. & Arn. (syn.: W. apiculata Sch.Bip.); CO, VE, EC, PE; 3600-4400 m W. villosa A.Gray; PE; 3300-4000 m Westoniel/a Cuatrec. Re f.:. Cuatrecasas, 1977. W. barqueroana Cuatrec.; CR; 3350-3800 m W. chirripoensis Cuatrec.; CR; 3400-3800 m W. eriocephala (Klatt) Cuatrec.; CR; 3100-3800 m W. kohkemperi Cuatrec.; CR; 3400-3800 m W. lanuginosa Cuatrec.; CR, PA; 3200-3350 m W. triunguifolia Cuatrec.; CR; 3350-3800 m Xenophyllum V.A.Funk Ref.: Funk, 1997a, 1997b. X crasswn (S.F.Biake) V.A.Funk (syn.: Werneria crassa S.F.Blake); CO, EC; 3650-4500 m X. fontii (Cuatrec.) V.A.Funk (syn.: Werneria fontii Cuatrec.); CO; 4200--4800 m X. humile (Kunth) V.A.Funk (syn.: Werneria articulata S.F.Biake, W. humilis Kunth); CO, EC, PE; 2800--4600 m X rigidum (Kunth) V.A.Funk (syn.: Werneria rígida Kunth); CO, EC; 4000--4800 m X. roseum (Hieron.) V.A.Funk (syn.: Werneria rosea Hieron.); EC; 3600--4300 m X sotarense (Hieron.) V.A.Funk (syn.: Werneria leucobryoides S.F.Biake, W. sotarensis Hieron.); CO, EC; 3800-4400 m
BALANOPHORACEAE Corynaea Hook.f. C. crassa Hook.f. var. sprucei (Eichl.) B.Hansen; CR, PE; 3350 m
BASELLACEAE Tournonia Moq. T. hookeriana Moq.; EC; 3500-3600 m
BEGONIACEAE Begonia L. [Note: Mainly a forest genus, but sorne species are found in subpáramo forest patches.J
[VOL. 84
B. difji1sa L.B.Sm. & B.G.Schub.; CO; 3500-3700 m B.ferruginea L.f.; CO; 2900--3300 m B. microphylla A.DC.; VE; 3100-3350 m B. montana (A.DC.) Warb.; VE; 3100-3400 m B. umbellata Kunth; CO; 3300-3600 m B. urticae L.f.; CO, EC, PE; 3100-3800 m; includes var. retusa L.B.Sm. & B.G.Schub. and-var. urticae
BERBERIDACEAE Berberis L. B. acutinervia L.A.Camargo; CO; 3250-3850 m B. aristeguietae L.A.Camargo; VE; 3600 m B. chimboensis C.K.Schneid.; CO; 3700-3900 m B. cretata L.A.Camargo; CO; 3000-3450 m B. cuatrecasasii L.A.Camargo; CO; 3450-3500 m B. densa Triana & Planch.; CO; 3200--4100 m B. discolor Turcz.; VE; 3200-3400 m B. dumaniana L.A.Camargo; CO; 3150-3200 m B.farinosa Benoist; EC; 3800 m B. goudotii Triana & Planch.; CO; 3000-3900 m B. grandiflora Turcz.; Cb, EC; 3000--4000 m B. cf. huertasii L.A.Camargo; CO; 2750-3900 m B. hyperythra Diels; EC; 3350-3450 m B.jamesonii Lindl.; EC; 3000--4000 m B. lutea Ruiz & Pav. var.lutea (syn.: B. conferta Kunth var. conferta and var. supranervata L.A.Camargo ); CO, EC, PE; 3050-3850 m B. meol/acensis L.A.Camargo; CO; 3250-3300 m B. minzaensis L.A.Camargo; EC; 3800 m B. nevadensis L.A.Camargo; CO; 3000-3100 m B. petriruizii L.A.Camargo; CO; 35SO m B. pichinchensis Turcz.; EC; 3000--4000 m B. prolifica Pittier; VE; 3100-3900 m B. psi/apoda Turcz.; CO?, VE; 3050-3475 m [Note: A species of limestone outcrops.) B. quinduensis Kunth; CO, EC; 3000-3650 m B. rigida Hieron.; CO, EC; 3300-3900 m B. rigidifolia Kunth; CO; 3025-3700 m B. stuebelii Hieron.; CO; 3050-3500 m B. sumapazana L.A.Camargo; CO; 3500-3800 m B. truxil/ensis Turcz.; VE; 3000-3900 m B. val/ensis L.A.Camargo; CO; 3450-3500 m B. verticillata Turcz.; CO; 3400-3750 m B. vitellina Hieron.; VE; 4100 m Mahonia Nutt. M. va/canica Stand!. & Steyer.; CR; 3100-3800 m
BIGNONlACEAE Eccremocarpus Ruiz & Pav. E. longiflorus Ruiz & Pav.; CO; 2700-3500 m
BORAGINACEAE Cynoglossum L. C. amabile Stapf & J.R.Drumm.; CR, EC; 3300-3500 m C. trianaeum Wedd.; CO; 3200 m
\
PARAMOS
1999]
)
Hackelia Opiz H. mexicana (Schltdl. & Cham.) l.M.Johnst.; CR; 20003500m H. revoluta (Ruiz & Pav.) I.M.Johnst. [syn.: H. cdstaricensis (Brand.) I.M.Johnst.]; CO, VE, EC; 3200-4400 m H. skutchii I.M.Johnst.; CR; 3500-3820 m Lappula Moench L. echinata Gi1ib.; CO; 3680 m [Note: Introduced from Europe.] Lithospermum L. L. columbianum Brandegee; VE; 2100-3000 m Moritzia DC. ex Meisn. M. lindenii (A.DC.) Benth. ex Gürke; CR, CO, VE, EC, PE?; 3250-4000 m Myosotis L. M. azorica R.C.Watson; VE; 2600-3300 m Plagiobothrys Fisch. & C.A.Mey. P. linifolius (Willd. ex Lehm.) I.M.Johnst.; CO, EC; 32003600 m . P. pygmaeus (Kunth) I.M.Johnst.; EC; 3200-3900 m Tournefortia L. T.fuliginosa Kunth; CO, EC; 3300-3550 m T. tarmensis (K.Krause) J.F.Macbr.; PE; 3100-3600 m
BRASSICACEAE Brassica L. B. integrifolia (West) O.E.Schulz; CO; 3700-3800 m Brayopsis Gilg & Muschl. B. colombiana AI-Shehbaz; CO, EC; 4300-4600 m; including subsp. colombiana and subsp. ecuadoriana Capsel/a Medik. C. bursa-pastoris (L.) Medik.; CO, EC, VE, PE; 2800-4000 m [Note: Introduced from Europe and cultivated.] Cardamine L. C. africana L.; CO; 3500-3700 m [l\ote: Introduced from Africa?] C. a/berti O.E.Schulz; CO; 3200-4100 m C. bonariensis Pers. (syn.: C. jlaccida Cham. & Schltdl.); CR, CO, VE, EC, PE; 3300-4500 m C. ecuadorensis Rieron.; EC; 3400 m C. hirsuta L.; CO, EC; 3550 m C.jamesonii Rook.; CO, EC; 3250-4200 m C. killipii O.E.Schulz; CO, VE; 3300-3800 m C. lehmannii Rieron.; CO, EC; 3450-3900 m C. lojanensis A 1-Shehbaz; EC; 3000-3450 m C. obliqua Rochst. ex A.Rich. (syn.: C. rhizomata Rollins); EC; ?m C. ovala Benth.; CR, CO; 2800-3350 m Cremolobus (Lam.) DC. C. peruvianus (Lam.) DC.; EC, PE; 2400-3600 m
99
Descurainia Webb & Berthel. D. myriophylla (Willd. ex DC.) R.E.Fries (syn.: Sisymbrium myriophyllum Willd. ex DC.); CO, EC, PE; 3500-4350 m Draba L. Ref.: Rangel Ch. & Santana C., 1989; Santana Castañeda, 1994. D. alyssoides Rumb. & Bonpl.; VE, EC, PE; 3700-4600 m D. arauquensis Santana; CO; 4360 m D. arbuscu/a Rook.f.; VE; 4000-4800 m D. aretioides Kunth (syn.: D. benthamiana Gilg); EC; 42004700 m D. barclayana AI-Shehbaz; CO; 4250-4400 m D. bellardii S.F.Biake; VE; 3500-4780 m D. boyacana AI-Shehbaz; CO; 4300-4550 m D. cheiranthoides Rook.f.; CO; 3400-5250 m D. chionophila S.F.Biake; VE; 4230-4750 m D. cocuyana AI-Shehbaz; CO; 3600-4050 m D. cryophila Cuatrec.; CO, VE; 4000-4500 m D. cuatrecasana Rangel & Santana; CO; 4000 m D. cuzcoensis O.E.Shultz; PE; ? m D. depressa Rook.f.; CO, EC, PE; 4200-4800 m D. extensa Wedd.; EC; 4500-4600 m D. farsetioides Linden & Planch.; VE; 4000-5200 m D.funckiana Linden & Planch.; VE; 4150-4500 m D.funckii (Turcz.) AI-Shehbaz; VE; 3100 m D. grandiflora Rook. & Arn.; EC; 3600-3800 m D. hallii Rook.f.; CO, EC; 3700-4800 m D. hammenii Cuatrec. & Cleef; CO; 4300-4600 m D. hookeri Walp.; EC; 3800-4300 m D.jorullensis Kunth; CR; 3500-3820 m D. lindenii (Rook.) Planch.; VE; 3500-4300 m D. litamo L.Uribe; CO; 4300-4800 m; includes subsp. glabra Santana and subsp. litamo D. matthioloides Gilg & O.E.Schulz subsp. saundersii AIShehbaz; PE; 4400 m D. obovata Benth.; EC; 3900-5000 m D. pachythyrsa Triana & Planch.; CO; 4300-4350 m D. pamplonensis Planch. & Linden; CO, VE; 4000-4300 m D. pennell-hazenii O.E.Schulz; CO; 4100-4350 m D. pseudocheiranthoides AI-Shehbaz; CO; 3470-4270 m D. pulvinata Turcz. (syn.: D. empetroides Gilg & O.E.Schulz); VE; 3600-4780 m; includes subsp. berryi and subsp. pulvinata D. pycnophylla Turcz.; CO, EC; 2415-4000 m D. rangelii Santana, ined.; CO; 3500-4300 m [Note: This species is not meant to be published herein.] D. ritacuvana Al-Shehbaz (syn.: D. cocuyensis Santana & Rangel); CO; 4300-4350 m D. rositae Santana & Rangel; CO; 3700-4450 m; includes subsp. boyacensis Santana & Rangel, subsp. rosita e, and subsp. sumapaensis Santana & Rangel D. sanctae-martae O.E.Schulz; CO; 3850-4300 m; includes var. leiocarpa O.E.Schulz and var. sanctae-martae D. schultzeii O.E.Schulz; CO; 4200 m D. schusteri O.E.Schulz; PE; 3400-3650 m D. sericea Santana & Rangel; CO; 3650-3900 m D. splendens Gilg; EC; 3600-4250 m D. spruceana Wedd.; EC; 4100-4300 m
1
( lOO
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
[VOL. 84
D. steyermarkii Al-Shehbaz (syn.: D. violacea DC. var. .féh,¡naniana O.E.Schulz); EC; 2700-4400 m D. venezuelana Al-Shehbaz; VE; 3000-3400 m D. wurdackii Al-Shehbaz; PE; 3350-3900 m
G. G. G. G.
Erysimum L. E. ghiesbreghtii Donn.Sm.; CR; 3200-3820 m
Pitcairnia L'Hér. [Note: This genus is found mostly in forest, but sorne species are in subpáramo forest patches.] P. meridensis Klotzsch ex Mez; VE; 2500-2800 m P. petraea L.B.Sm.; CO; 2300-3350 m P. pungens Kunth; CO, EC, PE; 2700-3800 m P. tympani L.B.Sm.; VE; 2800-2900 m
Eudema Humb. & Bonpl. E. colombiana Al-Shehbaz; CO, EC; 4250-4300 m; in eludes subsp. ecuadoriana Al-Shehbaz E. nubigena Humb. & Bonpl.; EC; 3800-4800 m; includes subsp. nubigena and subsp.'remyana (Wedd.) A1-Shehbaz E. rupestris Humb. & Bonpl.; EC; 4250-4550 m Halimolobos Tausch H. hispidula (DC.) O.E.Schulz var. hispidula; CO, EC;
2250-3300 m Lepidium L. L. abrotanifolium Turcz. (syn.: L.fraseri The11.); EC; 35004500m L. bipinnatifidum Desv.; CO, VE, EC, PE; 3100-3800 m L. ecuadoriense Thell.; EC; 2800-4400 m Nasturtium R.Br. (see Rorippa) . Romanschulzia O.E.Schulz R. apetala Rollins; CR; 3000-3500 m Rorippa Scop. R. nasturtium-aquaticum (L.) Hayek (syn.: Nasturtium officinale R.Br.); CO, EC, PE; 3050-3400 m [Note: Introduced from Europe.]
BROMELIACEAE Ref.: Gilmartin, 1972; Luther & Sieff, 1994; L. B. Smith & Downs, 1974, 1977, 1979. Greigia Regel [Note: This genus is mostly found in forest, but in páramo it may occur in open, boggy si tes.] G. columbiana L.B.Sm. var. columbiana: CO; 3800-4200 m G. danielii L.B.Sm.; CO; 2800-3500 m G. exserta L.B.Sm.; CO; 3200-3300 m G. mulfordii L.B.Sm.; CO; 3450-4050 m; includes var. macrantha L.B.Sm. and var. mulfordii G. nubigena L.B.Sm.; CO; 3350 m G. racinae L.B.Sm.; CO; 3300 m G. stenolepis L.B.Sm.; CO; 3000-3600 m G. vulcanica André; CO, EC; 3100-3750 m Guzmania Ruiz & Pav. [Note: This genus is found mostly in forest, but sorne species are in subpáramo forest patches.] G. candelabrum (André) André ex Mez; CO; 3100-3300 m G. conjinis L.B.Sm.; CO, VE; 2700-3600 m [Note: This species is found in calcareous areas.] G. danielii L.B.Sm.; CO; 2700-3100 m
lychnis L.B.Sm.; CO, EC; 3100-3400 m mosquerae (Wittm.) Mez; CO, EC; 3000-3600 m palustris (Wittm.) Mez; CO; 3500 m cf. variegata L.B.Sm.; PE; 3800 m
PuyaMolina P. adscendens L.B.Sm.; PE; 2800 m P. á/picola L.B.Sm.; CO; 3)50-3900 m P. ángulonis L.B.Sm.; PE; 3200-3950 m P ..angusta L.B.Sm.; PE; 3450 m P. antioquensis L.B.Sm. & Read; CO; 3100-3450 m P. aristiguietae L.B.Sm.; CO, VE; 3000-3550 m P. bicolor Mez; CO; 2850-3000 m P. boyacana Cuatrec.; CO; 3300-3550 m P. brachystachya (Bak.) Mez; CO; 3160 m P. cardonae L.B.Sm.; VE; 3100-3300 m P. clava-hercules Mez & Sodiro; CO, EC; 3350-4500 m P. cleefii L.B.Sm. & Read; CO; 3700 m P. compacta L.B.Sm.; EC; 2800-3800 m P. coriacea L.B.Sm.; PE; 3650 m P. cryptantha Cuatrec.; CO; 3000-3400 m P. cuatrecasasii L.B.Sm.; CO; 3400-3800 m P. dasylirioides Stand!.; CR; 3200-3600 m P. dichroa L.B.Sm. & Read; CO; 3750 m P. exuta L.B.Sm. & Read; CO; 3000 m P. fastuosa Mez; EC, PE; 3350-3900 m P. gigas André; CO; 3050 m q P. glaucovirens Mez; PE; 2400-3400 m P. glomerifera Mez & Sodiro; EC; 3600-4000 m P. goudotiana Mez; CO; 2600-3550 m P. grantii L.B.Sm.; CO, VE; 3000 m P. hamata L.B.Sm.; CO, EC, PE; 2850-4100 m P. horrida L.B.Sm. & Read; CO; 3450 m P. killipii Cuatrec.; CO, VE; 3000-3250 m P. linea/a Mez; CO; 2800-3700 m P. maculata L.B.Sm.; EC; 3200-3400 m P. medica L.B.Sm.; PE; 3800 m P. nítida Mez (syn.: P. killipii Cuatrec., P. nítida var. glabrior L.B.Sm. & Read); CO, VE; 2800-4000 m P. nivalis Baker; CO; 3250 m P. nutans L.B.Sm.; EC; 3000-4200 m P. occidentalis L.B.Sm.; CO; 3500-3600 m P. ochroleuca Betancur & Callejas; CO; 2700-3240 m P. pratensis L.B.Sm.; PE; 3200 m P. pygmaea L.B.Sm.; EC; 3150-3550 m P. ramonii L.B.Sm.; PE; 3700-4000 m P. ramosa L.B.Sm.; PE; 3200-3800 m "/ P. retrorsa Gilmartin; EC; 3200-3700 m P. roldanii Betancur & Callejas; CO; 2700-31 50 m 1 P. santanderensis Cuatrec.; CO; 3500 m
101
PARAMOS
1999]
P. santosii Cuatrec.; CO; 3000-3600 m; includes var. santosii and var. verdensis Cuatrec. P. trianae Baker; CO, VE; 3400-4100 m; includes var. ampliar L.B.Sm. & Read and var. trianae P. venezuelana L.B.Sm.; CO, VE; 3200-3950 m P. vestita André; CO, EC; 2750-3600 m P. westii L.B.Sm.; PE; 3400-3450 m Racinaea M.A.Spencer & L.B.Sm. R. tetrantha (Ruiz & Pav.) M.A.Spencer & L.B.Sm. (syn.: Tillandsia tetrantha Ruiz & Pav.)· CO VE· 300()....3800 m· includes var. aurantiaca (Griseb.),M.A.Spe~cer& L.B.Sm: and var. minia/a (André) M.A.Spencer & L.B.Sm. Tillandsia L. T. biflora Ruiz & Pav.; CO; 2730-3300 m T. clavigera Mez; CO; 3300 m T. compacta Griseb.; CO; 3000-3600 m T. complanata Benth.; CO, VE, EC; 3100-3600 m T. cuatrecasasii L.B.Sm.; CO; 3400-3550 m T. ionochroma André ex Mez; PE; 3200-3600 m T. orbicularis L.B.Sm.; CO, EC; 3300--4100 m T. riocreuxii André; CO; 2900-3300 m T. stenoura Harms; PE; 3500 m T. tetrantha (see Racinaea tetrantha) T. turneri Bak. var. turneri; CO, VE; 2950-3650 m
CALLITRICHACEAE Callitriche L. C. heteropoda Enge1m. ex Hegelm.; PE; 4100 m C. nubigena Fassett; CO, VE; 3500--4000 m C. cf. stagnalis Scop.; CO, VE; 3500--4100 m C. terrestris Raf. subsp. subsessilis (Fassett) Bacigalupo; CO; 4100m
CAMPANULACEAE Ref.: Jeppesen, 1981; Wimmer, 1957, 1968.
Centropogon C.Pres1 C. australis (E.Wimm.) Gleason; VE; 3000-3300 m C.je(Jtherstonei Gleason; PE; 3200-3400 m C.ferrugineus (L.f.) Gleason; CO, VE, EC; PE; 3100-3800 m C. intonsus Gleason; EC; 3200-3300 m C. e f. preslii E.Wimm.; EC; 2800-3500 m C. wil/denowianus (C.Presl) E.Wimm. subsp. cylindricus (G1eason) McVaugh; CO; 3400-3500 m
Hypsela C.Pres1 H. reniformis (Kunth) C.Pres1; CO, EC; 3500--4300 m Lobe/ia L. L. irasuensis Planch. & Oerst.; CR, PA; 3400-3900 m L. modesta Wedd.; CO, VE; 3250-3950 m L. tenera Kunth; CO, VE, EC, PE; 3200--4000 m
BRUNELLIACEAE Ref.: Cuatrecasas, 1970, 1985a.
Brunel/ia Ruiz & Pav. [Note: This genus of shrubs to small trees is mostly found in subpáramo forest patches and in the high Andean forest.] B. colombiana Cuatrec.; CO; 3300-3350 m B.faral/onensis Cuatrec.; CO; 3500--4000 m B. glabra Cuatrec.; CO; 2500-3000 m B. integrifolia Szyszyl.; CO, VE; 2400-2800 m B. pallida Cuatrec.; CO; 2800 m B. cf. putumayense Cuatrec.; CO; 2400-3100 m B. stuebelii Rieron.; CO; 2700-3100 m B. cf. trianae Cuatrec.; CO; 2700-3100 m
BUDDLEJACEAE Ref.: Norman, 1982, and in press.
Buddleja L. B. bu/lata Kunth (syn.: B. lindenii Benth.); CO, VE; 29003700 m
B. incana Ruiz & Pav.; CO, EC, PE; 2700--4500 m B.jamesonii Benth.; EC; 3000--4000 m B. nitida Benth. (syn.: B. alpina Oerst.); CR; 2500-3800 m B. pichinchensis Kunth; CO, EC; 3300--4300 m
CACTACEAE Opuntia Mili. O.jloccosa Salm-Dyck; PE; 4000 m [Note: See Sagástegui Alva, 1989).]
Lysipomia Kunth [Note: This genus is almost totally endemic to páramo.] Ref.: Ayers, 1997; Jeppesen, 1981; McVaugh, 1955; Van der Hammen & Cleef, 1978. L. acaulis Kunth; EC; 3900--4500 m L. aretioides Kunth; EC; 2900--4000 m L. bilineata McVaugh; EC; 3700-3800 m L. bourgoini Emst; VE; 3200--4300 m L. cae~pitosa T.J.Ayers; EC; 3000-3400 m L. crassomarginata (E.Wimm.) Jeppesen; EC; 3000-3400 m L. cuspidata McVaugh; EC: 3300-3800 m L. cylindrocarpa T.J.Ayers; EC; 3000-3400 m L. globularis E.Wimm.; PE; 3300-3500 m L. gracilis (E.Wimm.) E.Wimm.; PE; 3300-3400 m L. hirta E.Wimm.; PE;? m L. hutchisonii McVaugh; PE; 3000-3100 m L. laciniata A.DC.; CO, VE; 3350--4250 m; includes subsp. fissicalyx McVaugh, subsp. meridensis McVaugh, and subsp. microsperma McVaugh; not subsp. lacinia/a which is a puna taxon L. laricina E.Wimm.; EC; 3300-3400 m L. lehmannii Hieran. ex Zahlbr.; EC; 3000-3500 m L. montioides Kunth; CO, EC; 3800--4650 m L. multiflora McVaugh; EC, PE; 4000--4100 m L. muscoides Hook.f.; CO, EC; 3200-4200 m; inc1udes subsp. muscoides and subsp. simulans McVaugh L. oellgaardii Jeppesen; EC; 2850--4000 m L. rhizomata McVaugh; EC; 3400-3550 m L. sparrei Jeppesen; EC; 3100-3800 m L. speciosa T.J.Ayers; EC; 3000-3500 m
102
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
L. sphagnophila Griseb. ex Wedd.; CO, EC; 350b-4450 m; iD"cludes subsp. angelensis Jeppesen, subsp. minar McVaugh, and subsp. variabilis McVaugh L. subpeltata McVaugh; PE; 3100-3250 m L. tubulosa McVaugh; EC; 3700-4100 m L. vitreola McVaugh; EC; 3810-4500 m L. wurdackii McVaugh; PE; 2850-2900 m Rhizocephalum Wedd. (see Lysipomia) Siphocampylus Pohl S. benthamianus Wa1p.; CO; 3300-3900 m S. columnae (L.f.) G.Don; coi 3150-4100 m S. ellipticus (Willd.) Vatke (syn.: S. funckeanus P1anch.); CO; 2900-3600 m; includes var.funckeanus (P1anch.) E.Wimm. and var. peritornus (E.Wimm.) E. Wimm. S. giganteus (Cav.) G.Don; CO, EC; 3400-3700 m S.jelskii Zah1br.; EC, PE; 3400-3750 m S. cf. macropodoides Zahlbr.; PE; 3400 m S. cf. mirabilis E.Wimm.; CO; 3400 m S. odontosepalus Vatke; EC; 3400 m S. paramicola McVaugh; CO; 3500-3600 m S. retrorsus. (Willd.) Vatke; CO; 3300-3500 m S. sceptrum Decne.; VE; 3800-3900 m
CAPRIFOLIACEAE Viburnum L. V. triphyllum Benth.; CO; 3300-3700 m V. venustum C.V.Morton; PA; 3100-3300 m
CARYOPHYLLACEAE Arenaría L. A. dicranoides Kunth; VE, EC'; 3600-4600 m A.jamesoniana Rohrb.; EC; 4700 m A. lanuginosa (Michx.) Rohrb.; CR, PA?, CO, VE, EC; 1300-4100 m; includes subsp. lanuginosa and subsp. saxosa (A.Gray) Maguire A. moritziana Pax; VE; 3700 m A. musciformis P1anch. & Triana; CO, VE; 2900-4150 m A. muscoides Kunth; EC; 4600 m A. nítida (Bartl.) Rohrb.; PE; 4000 m A. parvifolia Benth.; EC, PE; 3550-4600 m A. quirosii Standl.; CR; 3500-3800 m A. radians Benth.; EC; 4000+ m A. reptans Hemsl.; CR; 3500-3800 m A. serpens Kunth; CO, EC, PE; 3550-4300 m A. cf. soratensis Rohrb.; CO, EC; 3000-4500 m A. tetragyna Willd. ex Schltdl.; EC; 3700-4500 m A. venezuelana Briq. (syn.: A. jahnii S.F.Blake); CO, VE; 3000-4500 m Cerastium L. C. alpinum L.; CO; 4400 m C. andinum Benth.; EC; 4600 m [Note: This species is perhaps equa1 to C. mollisimum.] C. arvense L.; CO, EC, PE; 3200-4100 m; includes var. arvense and var. arvensiforme (Wedd.) Rohrb. [Note: This species is native to Europe.]
[VQL. 84
C. candicans Wedd.; EC; 3800-450.0 m C. cephalanthum S.F.Blake; CO, VE; 3500-4300 m C. danguyi J.F.Macbr.; EC, PE; 3400-4300 m C.jloccosum Benth.; CO, EC; 4000-4800 m C. guatemalense Standl.; CR; 3500-3800 m C. imbricatum Kunth; CO, EC, PE; 3800-4500 m C. kunthii Brig. (syn.: C. glutinosum Kunth); CO, VE, EC; 3400-4500 m C. meridense Linden & Planch.; CO?, VE; 3250-3600 m C. mollisimum Poir.; CO, EC, PE; 3200-4600 m C. subspicatum Wedd.; CO, PE; 3400-4500 m C. trianae Briq.; CO, EC, PE; 3600-4500 m C. triviale Link (syn.: C. caespitosum Gilib.); CR, PA ?, CO, EC; 3450-4500 m C. cf.. tucumanense Pax; CO; 36.00-4050 m C. yiscosum L. (syn.: C. glomerata Thuill.); CR, PA, CO, VE; · 3500-4300 m [Note: This species is native to Europe.] C. vulgatum L.; CO, EC, PE; 3000-4000 m C. willdenowii Kunth; CO, EC; 3000-4000 m Colobanthus Bartl. C. quitensis (Kunth) Bartl.; CO, EC; 3850-4600 m Drymaria Wil!d. ex Roem. & Schult. D. cordata (L.) Willd. ex Roem. & Schult.; CO; 3400-3700 m, D. engleriana (Muschl.) Baehni & J.F.Macbr.; PE; 35003900m D.frutescens Mattf.; PE; 3650 m D. ovala Willd. ex Roem. &. Schult.; VE, EC; 3600-3900 m D. paramorum S.F.Blake; CO, VE; 2900-3700 m D. vi/losa Cham. & Schltdl. subsp. palustris (Cham. & Schltdl.) Duke; CO, VE; 2300-4000 m Paronychia Mili. P. bogotensis Triana & Planch.; CO; 2600-3800 m P. libertadiana Chaudhri; PE; 3200-3800 m
1
Sagina L. S. procumbens L.; CR, CO, EC; 3200-3800 m Scleranthus L. S. annuus L.; EC; 3400-3800 m [Note: This species is cosmopolitan weed.]
cy.
Si/ene L. S. gallica L.; EC; 3400-3800 m [Note: This species is native to Europe.] S. thysanodes Fenzl (syn.: Lychnis thysanodes Hook.f.); EC; 3500-4500 m Spergularia (Pers.) J.Presl & C.Presl S. spruceana R.Rossbach; EC; 3600 m Stellaria L. S. ciliata Vahl ex Pers.; CR, EC; 3000-3800 m S. cuspidata Willd. ex Schltdl.; CR, PA?, CO, EC, PE; 504200 m S. leptopetala Benth.; CO, EC; 2700-4750 m S. media (L.) Cirillo; CO, VE, EC, PE; 3000-3800 m
PARAMOS
1999]
S. recurvata Willd. ex Schltdl.; EC; 3250-4400 m S. serpyllifolia Willd. ex Schltdl.; CO, EC, PE; 2350-4000 m
CELASTRACEAE Maytenus Molina [Note: This genus is mostly found in forest, but the fol1owing species may occur in subpáramo forest patches.] M. novogranatensis Cuatrec.; CO; 3300-3550 m M. cf. prunifolia C.Presl; CO; ? m M. cf. trianae Briq.; CO;? m M. woodsonii Lundell (syn.: M. vulcanicola Stand!.); CR, PA; 2500-3820 m
CHLORANTHACEAE Hedyosmum Sw. [Note: Most species are found in forest, but sorne are found at the páramo/hum id 1ow forest ecotone (C1eef, pers. comm.).] Ref.: Todzia, 1988. H. colombiana Cuatrec.; CO; 3000-3500 m H. crenatum Occhioni; CO?, VE; 2850-3500 m H. cumbalense H.Karst. (syn.: H. granizo Cuatrec.); CO, EC; 2500-4300 m H. luteynii Todzia; CO; 3000-3550 m H. scabrum (Ruiz & Pav.) Solms (syn.: H. hirsutum H.B.K.); PE; 2800-3650 m H. toxicum Cuatrec.; CO; 3400-3450 m
CLETHRACEAE Clethra L. C.fagifolia Kunth; CO; 3350 m C. fimbriata Kunth; CO, EC; 3100-4000 m C. gelida Stand!.; CR; 3000-3500 m C. ovalifolia Turcz.; CO, EC; 2800-3600 m C. revoluta (Ruiz & Pav.) Spreng.; PE; 2850-3600 m
CLUSIACEAE Clusia L. [Note: Most species are found in forest, but sorne may occur in subpáramo thickets or at the páramo/humid 1ow forest ecotone (Cleef, pers. comm.).] C. alafa Triana & Planch.; CO; 3500-3700 m C. multiflora Kunth; CO; 3500-3900 m Hypericum L. Ref.: Robson 1987, 1990. H. aciculare Kunth; EC, PE; 3600-4000 m H. andinum G1eason; PE; 2400-4200 m H. baccharoides Cuatrec.; CO, VE; 2700-3350 m H. brathys (see H.juniperinum) H. brevistylum Choisy; EC, PE; 2400-4200 m H. bryoides G1eason; CO; 3300-4200 m H. ca/lacallanum N.Robson; PE; 3000-3750 m H. caracasanum Willd.; CR, VE; 3800-4100 m
103
H. cardonae Cuatrec.; CR, CO, VE; 2500-4250 m H. carinosum R.KeJler (syn.: H. stenoclados Cuatrec.); CO, VE; 2600-4500 m H. costaricensis N.Robson; CR, PA, CO; 2440-3700 m H. cymobrathys N.Robson; CO; 2970,.-3500 m H. decandrum Turcz.; EC; 3500-4200 m H. garciae Pierce; CO; 3100-3950 m H. gleasonii N.Robson; CO; 3200-4000 m H. gnidioides Seem.; CR, EC; 3200-3950 m H. goyanesii Cuatrec.; CO; 3000-3600 m H. horizontale N.Robson; CO; 2800-4500 m H. humboldtianum Steud. (syn.: H.jussiaei P1anch. & Linden); CO; 3000-3900 m H. irazuense Kuntze; CR, PA; 2700-3800 m H.jahnii (see H.juniperinum) H. jaramilloi N.Robson; CR, CO; 2600-3950 m H.juniperinum Kunth (syn.: H. brathys Sm., H.jahnii R.Keller, H. pseudobrathys Turcz.); CO, VE; 3100-3800 m H. lancifolium G1eason; CO, VE, PE; 2600-3900 m H. lancioides Cuatrec.; CO, VE, EC; 3100-4500 m; includes subsp. congestiflorum (Triana & P1anch.) N.Robson and subsp. lancioides H. laricifolium Juss. (syn.: H. laricoides G1eason); CO, VE, EC, PE; 3000-4300 m H. llanganatecium N.Robson; EC; 3250-4200 m H. loxense Benth.; EC, PE; 2350~3800 m; inc1udes subsp. aequatoriale (R.Keller) N.Robson and subsp. loxense H. lycopodioides Triana & Planch.; CO; 3350-4100 m H. magdalenicum N.Robson; CO; 2400-4875 m H. magniflorum Cuatrec.; CO, VE; 3100-3800 m H. marahuacanum N.Robson subsp. strictissimum N.Robson; CO, VE; 3000-3050 m H. martense N.Robson; CO; 3000-3900 m H. matangense N.Robson; EC; 3300 m H. mexicanum L.f.; CO, VE; 3100-4200 m H. myricariifolium Hieran.; CO; 3000-4100 m H. papillosum N.Robson; CO; 3550-3900 m H. parallelum N.Robson; CO; 3700-3950 m H. paramitanum N.Robson; VE; 2800-3150 m H. phellos G1eason (syn.: H. platyphyllum Gleason, H. tamanum Cuatrec.); CO, VE; 2500-4200 m; includes subsp. oroqueanum N.Robson, subsp. phellos, and subsp. platyphyllum (G1eason) N.Robson H. pimeleoides P1anch. & Linden ex Triana & P1anch. (syn.: H. caracasanum Willd. var. ocanense R.Keller); CO; 3100-3950 m . H. prostratum Cuatrec.; CO; 3200-4200 m H. quítense R.Keller; EC; 2600-3800 m H. recurvum N.Robson; PE; 2700-3300 m H. ruscoides Cuatrec.; CO, EC; 3300-3800 m H. sabiniforme Trevis.; CO; 3870 m H. selaginoides N.Robson; CO; 3300-4350 m H. simonsii N.Robson; CO; 3250-4100 m H. sprucei N.Robson; EC; 3500-4300 m H. stenopetalum Turcz. (syn.: H. meridense Steyerm.); CO, VE; 3050-4200 m H. strictum Kunth; CR, PA, CO, EC; 3100-4150 m; includes subsp. compactum (Triana & Planch.) N.Robson and subsp. strictum
104
H. H. H. H. H.
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
stue/;Jelii Hieron.; PE; 2400-3300 m tetrastichum Cuatrec.; CO, VE; 3700--4850 m thesiifolium Kunth; VE; 3900 m thuyoides Kunth; CO; 2500-3600 m va/leanum N.Robson; CO; 3750 m
Tillaea L. (see Crassula) Tillaeastrum Britton (see Crassula) Villadia Rose V. imbrica/a Rose; PE; 4000 m
COLUMELLIACEAE Columellia Ruiz & Pav. [Note: A forest genus that may occur at the forestpáramo ecotone.] C. lucida Danguy & Cherm.; EC; 390U m C. oblonga Ruiz & Pav. subsp. sericea (Kunth) Brizicky; CO; 3200-3600 m '
CONVOLVULACEAE Cuscuta L. C. grandijlora Kunth; CO, EC; 2700-3700 m Dichondra J.R.Forst. & G.Forst. D. microcalyx (Hallier f.) Fabris; PE; ? m
CORIARIACEAE Ref.: Skog, 1987.
Coriaria L. C. ruscifolia L. subsp. microphylla (Poir.) L.E.Skog (syn.: C. thymifolia Humb. & Bonpl. ex Willd.); CR, CO, VE, EC, PE; 3300-3500 m
CRASSULACEAE Crassula L. (syn.: Tillaea L.; Tillaeastrum Britton) Ref.: Bywater & Wickens, 1984. C. connata (Ruiz & Pav.) A.Berger; PE;? m [Note: det. A. Sagástegui A. Although the type is from Chancay (Peru), Bywater and Wickens (1984) say that it does not occur 1 in the Andes.] C. peduncularis (Sm.) Meigen [syn.: C. bonariensis (DC.) Cambess., C. paludosa (Schltdl.) Reiche, Tillaea paludosa S m., T. peduncularis Sch1tdl.]; CO, VE; 36004250 m C. venezuelensis (Steyerm.) M.Bywater & Wickens (syn.: Tilla ea venezuelensis Steyerm.); CO, VE, EC, PE; 28003950m C. viridis (S.Watson) M.Bywater & Wickens [syn.: Tillaea viridis S.Watson, Tillaeastrum viride (S.Watson) Britton] [Note: det. auct. Does not occur in Andes, according to Bywater and Wickens (1984).]
Echeveria DC. E. bicolor (Kunth) E.Wa1ther [syn.: E. bracteolata Link, K1otzsch & Otto, E. subspicata (Baker) A.Berger]; CO, VE, EC; 3200--4300 m
E. peruviana Meyen; PE; 2000-3500 m E. quitensis (Kunth) Lindl. [syn.: E. columbiana Poelln., E. sprucei (Baker) E.Morris]; CO, VE, EC; 3000-3900 m E. venezuelensis Rose; CO, VE; 3800--4100 m
[VOL. 84
CUNONIACEAE Weinmannia L. (Note: Primari1y a forest genus, but a1so found in shrub is1ands in páramo.] W. auriculata D.Don.; CO; 3100 m W. brachystachya Willd. ex Engl.; CO; 3000-3450 m W. elliptica Kunth; CO, EC; 3200-3600 m W. fagaroides Kunth; CR, CO, EC; 2700-4200 m (Note: ·This species is perhaps equa1 to W. microphylla.] W. glabra L.f. (syn.: W. caripensis Kunth); CR?, CO, VE, EC; 2000-3200 m [Note: This species is perhaps equa1 to W. pinnata L.] W. lansbergiana Engl.; CO; 3700-3900 m W. lopezana Cuatrec.; CO; 3300-3350 m W. mariquitae Szyszyl. (syn.: W. guanacasana Hieron., W. engleriana Hieron.); CO; 3200--4000 m W. microphylla Ruiz & Pav. (syn.: W. cochensis Hieron.); CO, EC, PE; 2800--4000 m; inc1udes var. microphylla and var. parvifolia Pamp. W. sibundoya Cuatrec.; CO; 2800 m W. silvatica Engl. (syn.: W. rollotii Killip); CO, PE; 31503200 m; inc1udes var. occidentalis Cuatrec. and var. rollotii (Killip) Cuatrec. W. cf. tamana Cuatrec.; CO; 2900 m W. tolimensis Cuatrec.; CO; 3400-3500 m W. tomentosa L. f.; CO; 2800--4000 m
CYPERACEAE Bulbostylis Kunth B. cf. glaziovii (Boeck.) C.B.C1arke; CO; 3000-3980 m [Note: Cleef 2236 (U) and other sheets at COL from 3000-3900 m are identified as B. tropicales (C.B.C1arke) Britton, a name that has apparent1y never been va1id1y pub1ished and that is not mean! to be va1idated herein.]
Carex L. (syn.: Vesicarex Steyerm.) C. acutata BÓott; CO, VE, EC; 3200--4000 m C. amicta Boott; CO, VE; 3000-3475 m C. cf. anisostachys Liebm.; CO, PE; 3300-3700 m C. azuayae Steyerm.; EC; 3800-3900 m C. boliviensis Van Heurck & Mü11.Arg.; PE; 3350--4100 m C. bonplandii Kunth s.l.; CR, CO, VE, EC, PE; 2700--4300 m C. brachyca!ama Boeck.; EC, PE; 3800--4100 m C. brehmeri Boeck.; CO; 3800--4100 m C. chordalis Liebm.; CR; 3100-3550 m C. co!lumanthus (Steyerm.) G.A.Whee1er (syn.: Vesicarex collumanthus Steyerm.); CO, VE; 3400--4300 m [Note: Mora's ( 1982) combination in Carex was inva1id; therefore, the combination mus! date from Wheeler' s (1989) paper, even though he rnade it inadvertent1y (T. Reznicek, pers. cornm.).]
'·
1999]
PARAMOS
C. crinalis Boott; CO, EC, PE; 2950-4300 m C. decidua Boott s.l.; CO, VE, PE; 3400-4000 m C. donnell-smithii L.H.Bailey; CR; 2700-3800 m C. ecuadorica Kük.; EC; 3960 m C. fecunda Steud. s.l.; CO?, PE; 3700 m C.jamesonii Boott; CR, CO, VE, EC, PE; 2800-4100 m C. lemanniana Boott; CR, PA, CO, EC; 2600-4000 m C. livida (Wah1enb.) Willd. (syn.: C. confertospicata Boeck.); CO, EC; 2500-4000 m C. luridiformis Mack. ex Reznicek & S.González; CO; 27003800m C. mandoniana Boeck.; EC, PE; 3000-4400 m C. microglochin Wahlenb. (syn.: C. oligantha Steud.); CO, EC, PE; 3500-4200 m C. aff. peucophila Ho!m; CO; 4050 m [Note: A close re1ative of C. mandoniana (T. Reznicek, pers. comm.).] C. phalaroides Kunth var. phalaroides; VE; 3050 m C. pichinchensis Kunth; CO, EC, PE; 3300-4350 m C. purdiei Boott; CR?, CO; 2700-3400 m C. pygmaea Boeck. (syn.: C. tristicha Spruce ex Boott non Boeck.); CR, PA, CO, EC; 3300-4500 m [Note: The o1dest name for C. tristicha may be C. tamana Steyerm. (T. Reznicek, pers. comm.), but the group needs more work.] i C. santae-marthae L.E.Mora & Rangel; CO; 3650 m C. setigluma Reznicek & S.González; CO, EC; 2800-3500 m C. tachirensis Steyerm.; VE; 3050-3450 m C. tamana Steyerm.; CR, CO, VE; 2700-4300 m C. toreadora Steyerm.; EC; 3800-4100 m Cyperus L. C. aggregatus (Willd.) ~ndl. [syn.: C. cayennensis (Lam.) Britton, C.jlavus (Vahl) Nees]; CO; 100-3950 m C. niger Ruiz & Pav.; EC; 3660 m C. rufus Kunth; CO; 2600-3500 m Eleocharis R.Br. E. acicularis (L.) Roem. & Schult. (syn.: E. nervata Svenson); CR, PA, CO, VE, EC; 2850-3800 m E. bonariensis Nees; EC; 3700 m E. capillacea Kunth; EC; 3300-3450 m E. dombeyana Kunth; CO, PE; 2800-4000 m E. exigua (Kunth) Roem. & Schult.; CO, VE; 3200-3750 m E. filiculmis Kunth; CO; 3580 m E. macrostachya Britton; CO; 2700-3850 m E. maculosa (Vahl) Roem. & Schult.; CO, PE; 3800-4300 m E. sellowiana Kunth; CO; 3800 m E. stenocarpa Svenson; CO, EC; 2800-4100 m
105
cleefii L.E.Mora; CO; 3600-4300 m ecuadorensis T.Koyama; CO, EC, PE; 3500-3800 m goeppingeri Suess.; CR, PA, CO, EC; 3000-4200 m obtusangulus Gaudich.; CO, EC, PE; 3300-4400 m; ineludes subsp. rubrovaginatus (Kük.) T.Koyama and subsp. unispicus Seberg O. venezuelensis Steyerm.; CR, PA, CO, EC; 2700-3800 m O. O. O. O.
Rhynchospora Vahl R. kunthii Nees ex Kunth; CR, CO, VE; 3200-3600 m R. macrochaeta Steud. ex Boeck.; CR, CO, VE, EC; 30004200m R. marisculus Nees ex Lindley & Nees; CO; 3000-3500 m R. oreoboloidea Gómez-Laur. (syn.: R. paramorum L.E.Mora non Steyerm.); CR, PA, CO; 3000-3550 m R. cf. paramora Steyerm.; VE; 2750-2950 m R. ruiziana Boeck.; CO, EC, PE; 3400-3800 m R. talamancensis Gómez-Laur. & W.W.Thomas; CR, PA, CO; 3000-3300 m R. tamana Steyerm.; VE; 3050-3450 m [Note: Species found on limestone outcrops.] R. tomentosa Steyerm.; VE; 3050-3450 m [Note: Species found on limestone outcrops.] R. vulcani Boeck.; CR, PA; 1200-3800 m Scirpus L. (see Jsolepis) Uncinia Pers. Ref.: Wheeler & Goetghebeur, 1997. U. ecuadorensis G.A.Whee1er; EC; 3900-4100 m U. koyamae Gómez-Laur.; CR; 3500-3820 m [Note: This species is perhaps equa1 to U. phleoides (Cav.) Pers. (G. A. Wheeler, pers. comm.).] U. lacustris G.A.Whee1er; EC; 3900-4100 m U. macrolepis Decne. (syn.: U. meridensis Steyerm.); CO, VE, EC, PE; 4000-4600 m U. paludosa G.A.Wheeler & Goetgh.; CO, EC, PE; 35004200m U. phleoides (Cav.) Pers.; CO, VE, EC; 3400-3950 m [Note: A species found primarily in montane forest (G. A. Wheeler, pers. comm.).] U. subsacculata G.A.Wheeler & Goetgh.; EC;3800 m [Note: A species found on1y in Polylepis forest (G. A. Wheeler, pers. comm.).] Vesicarex Steyerm. (see Carex)
DESFONTAINIACEAE
Jso/epis R.Br. (syn.: Scirpus L. p.p.) J. cernuua (Vahl) Roem. & Schu1t. (syn.: Scirpus cernuus Vahl); CO, EC, PE; 3600-4100 m l inundata R.Br. [syn.: Scirpus inundatus (R.Br.) Poir.]; CR, CO, VE, EC, PE; 3000-4200 m J. rígida Steud. [syn.: Scirpus rigidus (Steud.) Boeck.]; EC, PE; 3400-4100 m
Desfontainia Ruiz & Pav. Ref.: Leeuwenberg, 1969~ D. plowmanii R.E.Schu1t.; CO; 3500 m D. spinosa Ruiz & Pav. (syn.: D. parvifolia D.Don); CR, PA, CO, EC, PE; 2800-4000 m
Oreobolus R.Br. Ref.: Seberg, 1988.
Dioscorea L. D. larecajensis Uline ex R.Knuth; CO, PE; 2900-3600 m
DIOSCOREACEAE
106
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
DROSERACEAE Drosera L. D. cendeensis Tamayo & Croizat; VE; 2000-3000 m
ELAEOCARPACEAE Val/ea Mutis ex L. f.
[Note: A genus found mostly in forest, but sometimes also in distutbed páramo.] V. stipularis Mutis ex L. f.; CO, VE?, EC, PE; 3200-3800 m
ELATINACEAE Elatine L. E. chi/ensis Gay; CO; 3400'-4250 m E. ecuadoriensis Molau; CO, EC, PE; 2900'-4300 m E.fassetiana Steyerm.; CO, VE; 3500'-4000 m E. paramoana Schmidt-Mumm & R.Berna1; CO; 3000-
3700 m E. peruviana Baehni & J.F.Macbr.; PE;? m E. triandra Schkuhr; CO, VE, EC, PE; 2800-4000 m
EREMOLEPIDACEAE Antidaphne Poepp. & Endl. A. andina Kuijt; CO, EC; 1100-3350 m
ERICACEAE Ref.: Luteyn, 1995, 1996. Arctostaphylos Adans. (see Comarostaphylis) Bejaria Mutis ex L. ("Befaría")
Ref.: Clemants, 1995. B. aestuans L. [syn.: B. coarctata Bonpl., B. glauca Bonpl., B. oblonga (Ruiz & Pav.) Pers.]; CO, VE, EC, PE; 2600-
B. B. B.
B.
4000 m [Note: See Clemants, 1995, for a more extensive synonymy.] mathewsii Fielding & Gardner; CO, PE; 3100-3900 m nana A.C.Sm. & Ewan; CO, VE; 2900-3350 m resinosa Mutis ex L.f. (syn.: B. grandiflora Bonpl., B. weberbaueriana Mansf. & S1eumer); CO, VE, EC; 32004000 m [Note: See C1emants, 1995, for a more extensive synonymy.] tachirensis A.C.Sm.; CO, VE; 2900-3600 m
Cavendishia Lindl. C. bracteata (Ruiz & Pav. ex J.St.-Hil.) Hoerold [syn.: C. cordifolia (Kunth) Hoerold, C. crassifolia (Benth.) Hemsl., C. splendens (Klotzsch) Hoerold, C. strobilifera
(Kunth) Hoero1d]; CR, PA, CO?, VE, EC; 3100-4100 m [Note: See Luteyn, 1983, for a more extensive synonymy.] C. calycina A.C.Sm.; PA; 3100-3300 m C. ruiz-teranii Luteyn; VE; 3200-3300 .m Ceratostema Juss. C. a/atum (Hoero1d) S1eumer; CO, EC; 3250-4000 m
[VOL. 84
C. reginaldii (S1eumer) A.C.Sm.; EC; 2800-3340 m Comarostaphylis Zucc. C. arbutoides Lindl. subsp. arbutoides [syn.: Arctostaphy/os arbutoides (Lindl.) Hemsl.]; CR, PA; 2000-3820 m; inc1udes var. costaricensis (Small) Diggs Demosthenesia A.C.Sm. D. aff. microphyl/a (Hoero1d) A.C.Sm.; PE; 3600-3900 m
D. sp. 1; PE; 2800-3500 m [Note: An undescribed new species.] Disterigma (K1otzsch) Nied. D. acumina,tum (Kunth) Nied.; CO, EC; 2900-3800 m D. qlaterndides (Kunth) Nied.; CR, CO, VE, EC?, PE; 2500-
3800
rrl
D. codonanthum S.F.B1ake; EC; 3400-3800 m D. cuspidatum (P1anch. ex Wedd.) Nied.; CO; 2800 m D. empetrifolium (Kunth) Drude; CO, VE, EC, PE; 3150-
4300 m D. humboldtii (K1otzsch) Nied.; CR, PA, CO; 3100-3800 m D. microphyllum (G.Don) Luteyn; EC; 1800-3500 m D. staphelioides (P1anch.) Nied.; CO; 3100-3250 m Gau/theria L.
Ref.: Luteyn, 1995. G. amoena A.C.Sm.; CO, EC; 3050-4000 m G. anastomosans (L. f.) Kunth (syn.: G. pubiflora S.F.B1ake, G. ramosissima Benth.); CO, VE; 2800-4100 m G. bracteata (Cav.) G.Don; PE; 2650-3400 m G. buxifolia Willd. var. buxifolia; CO; VE; 2800-3300 m G. erecta Ven!. (syn.: G. cordifolia Kunth, G. loxensis Benth., G. meridensis A.C.Sm., G. odorata Kunth, G. ornata A.C.Sm., G. scabra Willd.); CR, PA?, CO, VE,
EC, PE; 3000-4000 m [Note: See Luteyn, 1995, for a more extensive synonymy.] G.fo/iolosa Benth.; CO, EC, PE; 3000-3800 m G. glomerata (Cav.) S1eumer; CO, VE, EC, PE; 33504150 m G. graci/is Small; CR; 1500-3100 m G. hapalotricha A.C.Sm.; CO, VE; 3200-4000 m G. insípida Benth.; CO, EC; 3300 m G. lanigera Hook.f. (syn.: G. rufo/anata S1eumer); CO, EC; 3200-3715 m; includes var.lanigera and var. rufo/anata (S1eumer) Luteyn G. megalodonta A.C.Sm.; EC; 3350-3700 m G. oreogena A.C.Sm.; CO, VE?, EC?; 3550-4300 m G. reticulata Kunth; EC; 3400-3900 m G. rígida Kunth (syn.: G. psilantha A.C.Sm.); CO, VE; 2900-3600 m G. santanderensis A.C.Sm.; CO; 3000-4100 m G. sclerophylla Cuatrec.; CO, VE, EC; 3300-4000 m; ineludes var. hirsuta Luteyn and var. sclerophylla G. strigosa Benth. (syn.: G. arachnoidea A.C.Sm.); CO, VE, EC; 2800-3950 m; inc1udes var. revoluta (A.C.Sm.) Luteyn and var. strigosa G. vaccinioides Wedd.; PE; 3000-3380 m
1
1999]
107
PARAMOS
Gaylussacia Kunth G. buxifolia Kunth; CO, VE; 2500-3500 m Mac/eania Hook. M. hirtijlora (Benth.) A.C.Sm.; CO, VE, EC; 3000-3715 m M. rupestris (Kunth) A.C.Sm.; CR, CO, VE, EC; 3250--4100 m [Note: See Luteyn, 1997, for extensive synonymy.] Pernettya Gaudich. Ref.: Luteyn, 1995. P. hirta (Willd.) Sleumer (syn.: Gaultheria purpurascens Kunth); CO; 3150-3700 m P. prostrata (Cav.) DC. [syn.: P. coriacea K1otzsch, P. myrsinoides (Kunth) Zucc. ex Steud., P. parvifolia Benth., Gau/theria myrsinoides Kunth]; CR, PA, CO, VE, EC, PE; 3300--4650 m [Note: See Luteyn, 1995, for a more extensive synonymy.] Plutarchia A.C.Sm. P. angula/a A.C.Sm.; CO, EC; 3150-3300 m P. coronaria (Lindl.) A.C.Sm.; CO; 2850-3750 m P. dasyphylla A.C.Sm.; CO; 2800-3600 m ¡P. ecuadorensis Luteyn; EC; 2960-3600 m P. guascense (Cuatrec.) A.C.Sm.; CO; 3400-3600 m P. minar A.C.Sm.; CO; 3400-3700 m ¡ P. miranda A.C.Sm.; CO; 2160-3400 m P. cf. monantha A.C.Sm.; CO; 3350 m P. pubijlora A.C.Sm.; CO; 3600-3700 m P. rigida (Benth.) A.C.Sm.; CO; 3300-3700 m Psammisia Klotzsch P. graebneriana Hoerold; CO; 3300 m P. hookeriana Klotzsch; VE; 3200 m P. /ehmannii Hoerold; CO; 3200-3300 m P. pendulijlora (Dunal) Klotzsch; VE; 900-3750 m Semiramisia Klotzsch S. pulcherrima A.C.Sm.; CO; 2890-3400 m
S. speciosa (Benth.) Klotzsch; CO; 1900-3140 m Themistoclesia Klotzsch T anfracta (A.C.Sm.) Sleumer; CO; 3200-3450 m T. campii A.C.Sm.; EC; 3200-3300 m T compta A.C.Sm.; CO; 3500-3600 m T. dependens ()3enth.) A.C.Sm. (syn.: T pendula K1otzsch); CO, VE; 1500-3400 m T dryanderae Sleumer; CO; 3500-3600 m T epiphytica A.C.Sm.; CO, EC; 3150-3500 m T mucronata (Benth.) Sleumer; CO; 3050-3800 m T vegasana A.C.Sm.; CO; 2900-3400 m
V. corymbodendron Dunal; CO, VE; 2000-3500 m V. crenatum (G.Don) Sleumer; EC; 2500-3700 m V.jloribundum Kunth (syn.: V. mortinia Benth.); CR, CO, VE, EC, PE; 3000-4350 m; includes var. marginatum (Dunal) Sleumer and var. ramosissimum (Duna!) S1eumer V. meridionale Swartz (syn.: V. caracasanum Kunth); CO, VE; 2000-3500 m V. poasanum Donn.Sm.; CR; 2000-3100 m
ERIOCAULACEAE Eriocaulon L. E. microcephalum Kunth; CO, EC, PE; 3350--4100 m Paepalanthus Kunth P. a/pinus Kiirn.; CO; 3200-3500 m P. andicola Kiim.; CO, VE; 3050-3750 m; includes var. andico/a and var. villosus Moldenke P. columbiensis Ruhland; CO, VE; 2500-3500 m P. costaricensis Moldenke; CR; 3000-3500 m P. crassicaulis Kiirn.; CO; 3000-3400 m P. diffisus Mo1denke; VE; 3200-3600 m P. ensifolius (Kunth) Kunth; CO, VE, EC; 3100-3300 m P. espinosianus Moldenke; EC, PE; 3450 m P. granatensis Kiirn.; CO; 2800 m P. karstenii Ruhland; CO, VE, EC, PE; 3100-4300 m; ineludes var. corei Moldenke, var. minimus Moldenke, and var. subsessilis (Moldenke) Moldenke P. kupperi Suess.; CR, PA; 3300-3600 m P. linde~ii Ruhland; CO; 2700-3600 m P. lodiculoides Moldenke (syn.: P. po/ytrichoides Kunth var. densus Moldenke; Syngonanthus steyermarkii Moldenke); CO, VE; 3700--4400 m; includes var.jloccosus Moldenke and var. lodiculoidr:s P. /oxensis Moldenke; EC; 2500-3500 m P. macarenensis Moldenke; CO; 3000-3100 m P. meridensis Klotzsch; CO, VE; 2900-3300 m P. muscosus Kiirn.; CO, VE; 2900-3400 m; includes var. muscosus and var. tachirensis Moldenke P. o/igocephalus Kiirn.; CO; 3250 m P. paramensis Moldenke; CO, PE; 3100-3600 m P. petraeus Kiirn.; CO; 3200-3400 m P. pi/osus (Kunth) Kunth; CR, CO, VE, EC, PE; 3000--4000 m P. planifolius (Bong.) Kiirn. var. a/pestris Kiirn.; CO; 26003650 m P. schlimii Kiirn.; CO; 2800-3200 m P. shultesii Moldenke; CO; 3000-3500 m P. stuebelianus Ruhland; PE; 2900-3250 m P. truxi/lensis Kiirn.; VE; 2500-3500 m [Note: This species is found on 1imestone outcrops.]
Thibaudia Ruiz & Pav. ex J.St.-Hil. T.fallax A.C.Sm.; CO; 3400-3500 m Tfloribunda Kunth; CO, EC?; 3000-3600 m T. grantii A.C.Sm.; CO; 3350 m T parvifolia (Benth.) Hoerold; CO, EC; 3000-3750 m
S. peruvianus Ruhland; PE; 2800-3100 m
Vaccinium L. V. consanguineum Klotzsch; CR, PA; 3100-3500 m
Chamaesyce Raf. C. ovalifolia Engelmann; CO; 3700 m
Syngonanthus Ruh1and
EUPHORBIACEAE
108
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
Dysopsis Baill. D. glechomoides (A.Rich.) Müll.Arg.; CR, CO?, EC; 36003750m Euphorbia L. E. coarctata (Ruiz & Pav.) Baker; CO, VE?, PE; 28003450 m
FABACEAE Astragalus L. A. garbanzillo Cav.; PE; 3000--4000 m A. geminiflorus Humb. & Bonpl.; EC; 4000-5000 m A. weberbaueri U1br.; PE; 2400-3700 m Cologania Kunth C. broussonetii (Ba1b.) DC.; CO; 2800 m Da lea L. [Note: These three species of Da/ea are from very weedy sites and are probab1y not true páramo species.] D. coerulea (L.f.) Schinz & Thell.; CO, EC; 1000--4000 m D. exilis DC.; PE; 2500-3700 m D. humifusa Benth.; EC; 1300-3100 m Lathyrus L. L. magellanicus Lam. (syn.: L. longipes Phi!.); CO, EC, PE; 3100--4000 m L. meridensis Pittier; CO, VE; 3000-3350 m Lupinus L. [Note: The genus Lupinus is current1y under study by Rupert Bameby (NY), who thinks that most determinations are uncertain and that there may only be about 15 true Andean species (pers. comm.).] L. alibicolor C.P.Sm.; PE; 3465 m L. alirevolutus C.P.Sm.; CO; 3150-3450 m L. alopecuroides Desr.; CO, VE, EC, PE?; 3450--4700 m L. argurocalyx C.P.Sm.; CO; 3200-3250 m L. arvensis Benth.; EC; 2900 m L. asymbepus C.P.Sm.; PE; 2900-3600 m L. austrohumifusus C.P.Sm. (syn.: L. humifusus Benth. non Sessé & Mo~.); CO, VE, EC, PE; 3300--4500 m L. bogotensis Benth. (syn.: L. amandus C.P.Sm., L. bogotensis var. parvior C.P.Sm.); CO, EC; 2800--4000 m L. boyacensis C.P.Sm.; CO; 3400-3500 m L. brevecuneus C.P.Sm.; EC; 3500 m L. carrikeri C.P.Sm.; CO; 3850 m L. chipaquensis C.P.Sm.; CO; 3150 m L. chrysocalyx C.P.Sm.; CO; 3950--4150 m L. co/ombiensis C.P.Sm.; CO; 4100--4400 m L. cotopaxiensis C.P.Sm.; EC; 3660 m L. eremonomus C.P.Sm.; VE; 4000 m L.falsorevolutus C.P.Sm.; CO; 3500 m L.famelicus C.P.Sm.; CO; 3800--4200 m L.fieldii Rose ex J.F.Macbr.; PE; 3500-3700 m L.foliolosus Benth.; EC; 3600--4500 m L. guascensis C.P.Sm.; CO; 3300-3700 m
[VOL. 84
L. interruptus Benth.; CO; 3900--4100 m L.jahnii Rose ex C.P.Sm.; VE; 3800--4000 m L. kunthii J.Agardh.; EC; 3400--4000 m L. lagunae-negrae C.P.Sm.; CO; 3700-3850 m L. lespedezioides C.P.Sm.; CO; 3400-3700 m L. cf. lindenianus C.P.Sm.; VE; 3600--4400 m L. lindleyanus J .Agardh; EC, PE; 2100--4000 m L. magdalenensis C.P.Sm. var. seifrizianus C.P.Sm.; CO; 3400 m 'L. meridanus Moritz ex C.P.Sm.; VE; 2000--4000 m L. michelianus C.P.Sm.; EC; 3500--4000 m L. microphyllus Desv.; CO, EC; 37~0--4600 m L. mirabilis C.P.Sm.; CO; 3400 m L. monserratensis C.P.Sm.; CO; 3000-3300 m L. mutabilis Sweet; CO, VE, EC, PE; 3800--4000 m [Note: .This species is often cultivated as a crop.] L. ~ubigenus Kunth; EC; 4100--4 700 m L. pachanoanus C.P.Sm.; EC; 3700 m L. panicu/atus Desr.; VE, EC; 3600 m L. patulus C.P.Sm.; EC; 3800--4200 m L. penlandianús C.P.Sm.; EC; 4200 m L. peruvi~nus Ulbr. (syn.: L. decemplex C.P.Sm., L. spragueanus C.P.Sm.); CO, VE, EC, PE; 3100--4150 m; includes var. spragueianus C.P.Sm. L. pseudotsugoides C.P.Sm.; EC; 4200 m L. puracensis C.P.Sm.; CO; 3500 m L. revolutus C.P.Sm.; EC; 4000 m L. ruizensis C.P.Sm.; CO; 3900--4200 m L. rupestris Kunth (syn.: L. caespitosus Benth. non Nuttall ex Torr. & A.Gray, 1840); EC; 4080--4100 m L. smithianus Kunth; EC; 4500 m L. subcuneatus C.P.Sm.; CO; 3200 m L. subhamatus C.P.Sm.; CO; 3820 m L. tamayoanus C.P.Sm.; VE; 4000 m L. tauris Benth.; EC; 3350-3450 m L. tolimensis C.P.Sm.; CO; 3100-3500 m L. trianus C.P.Sm.; CO; 3400 m L. varicaulis C.P.Sm.: CO; 3500--4200 m L. venezuelensis C.P.Sm.; VE; 3800--4200 m L. weberbaueri Ulbr.; PE; 3800-3900 m Medicago L. [Note: Species ofthis genus are native to Europe, introduced, and often cultivated in páramo regions as forage.] M. arabica (L.) Huds.; CO; 3300-3400 m M. polymorpha L.; VE; 2900-3400 m Otholobium C.H.Stirt. (syn.: Psoralea L. p.p.) O. caliginis J.W.Grimes; CO; 3300 m O. mexicanum (L. f.) J.W.Grimes [syn.: Psora/ea mexicana (L.f.) Vai1]; CO, EC, PE; 2800-3250 m
O. pubescens (Poir.) J.W.Grimes; PE; 3300 m Psoralea L. (see Otholobium) Trifolium L. T. amabile Kunth; CR, CO, EC, PE; 3220-3800 m [Note: Native weed used as forage.]
1
.¡
1999]
PARAMOS
T. dubium Sibth.; CR, CO; 3000-3200 m [Note: Native weed used as forage.] T. filiforme L.; CO, EC; 3450 m [Note: Native weed used as forage.] T. repens L.; CR, CO, EC; 3450-3800 m [Note: Native to Europe, introduced and cultivated as forage.]
Vicia L. [Note: Species of this genus are used as forage.] V. andicola Kunth; CO, VE, EC, PE; 3000-4100 m V. gramínea Sm.; CO, EC, PE; 3150-3500 m
GARRYACEAE Gan·ya Douglas ex Lindl. G. laurifolia Hartweg ex Benth. subsp. quichensis Donn.Sm.; CR; 3250-3800 m
GENTIANACEAE Ref.: Ba1slev & Briones, 1982; Pringle, 1995.
Gentiana L. G. arbelaezii Cuatrec.; CO; 3200 m G. cocuyana Cuatrec.; CO; 3350-4300 m G. sedifo/ia Kunth (syn.: G. pros trata Haenke); CR, PA, CO, VE, EC, PE; 2800-4 700 m Gentianella Moench [Note: Essentially a páramo genus.] Ref.: Fabris, 1958, 1960. G. amoena (Wedd.) Fabris; PE; 3500-3820 m G. androsacea J.S.Pringle; EC; 3100-3600 m ' G. atroviolacea (Gilg) J.S.Pringle (syn.:. Gentiana atroviolacea Gi1g); CO; 3500 m G. bicolor (Wedd.) J.S.Pringle; PE; 3400-3550 m G. brunneotincta (Gi1g) J.S.Pringle (syn.: Gentiana brunneotincta Gilg); PE; 3800-4100 m G. cerastioides (Kunth) Fabris; CO, EC, PE; 2800-4700 m G. cernua (Kunth) Fabris; EC; 3300-5000 m G, chamuchui (Reimers) Fabris; PE; 3700-4100 m G. corallina (Gi1g) Zarucchi; PE; 3500-3700 m G., cor)mzbosa (Kunth) Weaver & Rüdenberg; CO, VE; 3200-3900 m G. crassicaulis (Gilg) J.S.Pringle (syn.: Gentiana crassicaulis Gilg); PE; 3750 m G. crassulifolia (Griseb.) Fabris [syn.: Gentiana crassulifolia Griseb., Gentianella lehmannii (Gilg) Fabris]; EC; 32004100m G. dacrydioides (Gilg) Weaver & Rüdenberg [syn.: Gentiana dacrydioides Gilg; Gentianella eng/eri (Gilg) Fabris]; CO, EC; 3250-3600 m G. dasyantha (Gilg) Fabris; CO; 3700-4400 m G. dianthoides (Kunth) Fabris ex J.S.Pringle (syn.: Gen liana dianthoides Kunth); CO, PE; 3750-4200 m G.fastigiata Fabris; EC; 3000-3800 m G.flaviflora (Gi1g) Fabris (syn.: Gentianajlaviflora Gilg); EC; 4500-4550 m G.foliosa (Kunth) Fabris (syn.: Gen liana androti"icha Gilg);
109
EC; 3200-4800m G.fuscicaulis Fabris; EC; 3100-3650 m G. gilgiana (Reimers) Fabris ex J.S.Pringle (syn.: Gentiana gilgiana Reimers); PE; 4000-4700 m G. gilioides (Gi1g) Fabris; EC; 2700-3500 m G. gracilis (Kunth) Fabris; EC; 3000-3300 m G. gramínea (Kunth) Fabris (syn.: Gentiana gramínea Kunth); PE; 3200-3750 m G. hirculus (Griseb.) Fabris; EC; 3650-4400 m G. hypericoides (Gilg) Fabris; EC; 3000-3500 m G. hyssopifolia (Kunth) Fabris; EC; 2700-3800 m G.jamesonii (Hook.) Fabris; EC; 2850-4150 m G. limoselloides (Kunth) Fabris; EC; 2700-4700 m G. longibarbata (Gilg) Fabris; EC; 3300-4400 m G. nevadensis (Gilg) Weaver & Rüdenberg (syn.: Gen tia na nevadensis Giig); CO, VE, EC; 3300-4250 m G. nummulariifolia (Griseb.) Fabris; CO, EC; 3300--4650 m G. oelgaardii J.S.Pringle; EC; 3000-3700 m G. oreosilene (Gilg) J.S.Pringle; PE; 3400-3750 m G. pernettyoides (Reimers) Fabris; PE; 3450--4000 m G. rapunculoides (Willd. ex Schult.) J.S.Pringle [syn.: G. diffusa (Kunth) Fabris, G. hookeri (Griseb.) Fabris]; CO, EC; 3200-4300 m G. rubra J.S.Pringle, ined.; PE; 3900 m [Note: This species is not mean! to be published herein.] G. rugicalyx J.S.Pringle, ined.; PE; 4000 m [Note: This species is not mean! to be published herein.] G. rupicola (Kunth) Holub (syn.: Gen liana rupicola Kunth); EC; 3750-4600 m G. sagasteguii J.S.Pringle, ined.; PE; 3600-4000 m [Note: This species is not meant lo be published herein.] G. saxifragoides (Kunth) Fabris; EC; 2700-3600 m G. selaginifolia (Gilg) Fabris (syn.: Gentiana solidagoides Reimers); CO; 3400--4500 m G. stellarioides (Griseb.) Fabris; CO; 3300-3850 m G. sulphurea (Gilg) Fabris; EC; 4000--4500 m G. thyrsoidea (Hook.) Fabris; PE; 4000 m G. tristicha (Gilg) J.S.Pringle; PE; 3900 m G. uberula J.S.Pring1e, ined.; PE; 3300--4000 m [Note: This species is not meant to be published herein.] G. vaginalis (Griseb.) J.S.Pringle; PE; 3900 m G. viridis (Griseb.) Weaver & Rüdenberg (syn.: Gentiana viridis Griseb.); VE; ? m
Halenia Borkh. Ref.: Allen, 1933. H. adpressa C.K.Allen; CO; 3150-3900 m H. aquilegiella Stand!.; CR; 3000-3800 m H. asclepiadea (Kunth) G.Don; CO; 3300-3750 m H. campanulata Cuatrec.; CO; 4000 m H. cuatrecasasii C.K.Allen; CO; 2800-3300 m H. dasyantha Gilg; CO; 4000-4875 m H. elegans C.K.Allen; CO; 3800-4100 m H. foliosa Gilg; CO; 2800--4000 m H. garcia-barrigae C.K.Allen; CO; 3200 m H. gentianoides Wedd.; CO, VE; 3500--4350 m H. gigantea C.K.Allen; CO, VE; 3500--4150 m H. gracilis (Kunth) G.Don; CO; 3500-3900 m
\ 110
MEMOIRS OF THE NEW YORK BOTANICAL O ARDEN
H .h<Jppii Reimers; CO; 3800-4300 m H hygrophila Gi1g; CO; 3000-4300 m H hypericoides (Kunth) G.Don; CO; 3050 m H inaequalis Wedd.; VE; 3000-4300 m H insignis C.K.Allen; CO; 3800-4500 m H kalbreyeri Gilg; CO, EC; 2900-4100 m H karstenii Gi1g; CO; 3600-3700 m H. longicaulis J.S.Pring1e; EC; 2400-3800 m H macrantha Gi1g; CO; 3550-3700 m H. majar Wedd.; CO; 3300-4000 m H mínima C.K.Allen; EC; 2500-3900 m H pauana Cuatrec.; CO; 3300-4050 m H. phy/lophora C.K.Allen; CO; 2900-3250 m H pu/chella Gi!g; EC; 3300-4400 m H. purdieana Wedd. var. congesta C.K.Allen; CO; 3000-c 4300m H rhyacophila C.K.Allen; CR; 2500-3800 m H schultzei Gilg ex C.K.Allen; CO; 3300-3850 m H serpyl/ifolia J.S.Pring1e; EC; 3000-4100 m H. stellarioides Gi1g; CO; ? m H. subinvolucrata Gi1g; VE; 1300-4900 m H. taruga-gasso Gi1g; EC; 2800-4200 m . H. tolimae Gi1g; CO; 3900-4500 m H. umbellata (Ruiz & Pav.) Gilg; PE; 3600 m H. venezue/ensis C.K.Allen; VE; 3300-4000 m H. verticil/ata Gi!g; CO; 3500-3700 m H. viridis (Griseb.) Gilg; VE; 3300-4300 m H. weddelliana Gi1g (syn.: H. meyer-johannis Gilg); CO, EC; 3300-5000 m Macrocarpaea (Griseb.) Gi1g M. glabra (L. f.) Gilg; CO; 2800-4000 m M pachyphylla Gilg; CO, EC; 3250-3800 m M. stenophylla Gi!g; PE; 2850-3300 m
GERANIACEAE Erodium L'Hér. ex Aiton E. cicutarium (L'Hér.) ex Aiton; CR, VE, EC; 3300-3700 m [Note: Intraduced fram Europe and used medicinally.] E. moschatum (L'Hér.) ex Aiton; CO, VE; 3000-3900 m [Note: Introduced fram Europe.] Geranium L. Ref.: Ha1fdan-Nielsen, 1995. G. agavacense Willd.; EC; 2400-3250 m [Note: Based on the Bonpland collection, type photo F neg. 35705.] G. ange/ense Halfdan-Niels.; CO, EC; 3700-4500 m (5, antisanae R.Knuth; EC; 4050-4200 m G. azorel/oides Sandwith (syn.: G. guanacosense R.Knuth); CO, EC; 3200-3450 m G. campii H.E.Moore; EC, PE; 3000-3650 m G. chilloense Willd. ex Kunth (syn.: G. schimpffii R.Knuth); CO, EC, PE; 2230-3600 m G. chimborazense R.Knuth; EC; 3580-4300 m G. co/umbianum R.Knuth; CO; 3200-3500 m G. costaricense H.E.Moore (syn.: G. cucul/atum Kunth var. multifidum Suess., G. bolivarianum W.A.Dayton); CR, PA; 2950-3800 m
[VOL. 84
G. diffusum Kunth (syn.: G. quinque/obum Wedd., G. imbaburae R.Knuth); CO, VE, EC, PE; 2300-4000 m G. dissectum L.; CO; 3500 m [Note: This species is native to Europe and Asia, now naturalized in páramo regions.] G. ecuadoriense Hieran.; EC; 4100-4650 m G. exal/um H.E.Moore (syn.: G. jahnii Stand!.); VE, 'Eé; 3050-3400 m G. guamanense Halfdan-Nie1s.; EC; 3950-4100 m G. hirtum Willd. ex Spreng.; CO, VE; 2850-3700 m G. holm-nielsenii Halfdan-Niels.; EC; 4-200 m G. humboldtii Willd. ex Spreng. (syn.: G. acau/e Willd. ex Kunth); EC; 3600-4500 m G. /axicaule R.Knuth; CO, EC, PE; 2500-3800 m G. lignosum R.Knuth; CO; 3200 m G. lindenianum Turcz.; CO; 3650 m G.. 'loxense Halfdan-Niels.; EC; 2500-3450 m G. maniculatum H.E.Moore; CO, EC; 3300-5000 m G.. meridense Pittier; VE; 3100 m G. mogotocorense R.Knutll; CO; 3700--:3800 m G. multiceps Turcz.; CO, VE; 3300-4200 m G. multipartitum Benth. (syn.: G. heinrichsae R.Knuth); CO, EC, PE; 3250-4700 m; includes var. glabrescens Hieran. ex R.Knuth, var. multipartitum, and var. velutinum R.Knuth G. paramicola R.Knuth; CO; 3850-3900 m G. pavonianum Briq.; PE; 3500 m G. peruvianum Hieran.; PE; 3300-4000 m G. repens H.E.Moore; CR, PA; 3000-3400 m G. reptans R.Knuth (syn.: G. caucense R.Knuth); CO, EC, PE; 2900-4600 m G. santanderiensis R.Knuth; CO; 3300-4100 m G. schimpfii Kunth; CO; 3000-3300 m G. schultzei R.Knuth; CO; 3000-3900 m G. sebosum S.F.Blake; VE; 4000 m G. sericeum Willd. ex Spreng.; EC; 3900-4600 m G. sibbaldioides Benth. (syn.: G. cucul/atum Kunth, G. igualatense R.Knuth); CO, VE, EC, PE; 2950-4400 m; includes var. e/ongatum Wedd. and var. sibbaldioides G. stoloniferum Stand!.; VE; 3000 m G. stramineum Triana & Planch. (syn.: G. confertum Stand!.); CO, EC; 3000-4100 m G. subnudicau/e Turcz.; CO, VE; 3750 m G. tracyi Sandwith; VE; 2900 m G. velutinum Turcz.; CO, VE; 2860-3900 m G. venezue/ae R.Knuth; VE; 3300-3750 m
GESNERIACEAE
1
·1
Al/oplectus Mart. [Note: A forest genus, but rarely found in subpáramo forest patches.] A. ichthyoderma Hanst.; CO, VE; 2300-3750 m A. peruvianus (Zahlb.) L.P.Kvist &· L.E.Skog; CO; 33003350 m Co/umnea L. [Note: A forest genus, but rarely found in subpáramo forest patches.] C. strigosa Benth.; CO, VE; 3200-3600 m
1999]
111
PARAMOS
Heppiella Rege1 H. ulmifolia (Kunth) Hanst.; CO, EC; 2400-3800 m
GROSSULARIACEAE Escallonia Mutis ex L. f. E. myrtilloides L. f. [syn.: E. cmymbosa (Ruiz & Pav.) Pers., E. poasana Donn.Sm., E. tortuosa Kunth]; CR, PA, CO, VE, EC, PE; 1700-4100 m; in eludes var. myrtilloides and var. patens (Ruiz & Pav.) S1eumer Ribes L. R. andicola Jancz.; CO, EC; 3100-4300 m R. bogotanum Jancz.; CO; 3200-3350 m R. canescens Pittier; VE; 3600-3650 m R. ciliatum Humb. & Bonpl. ex Roem. & Schu1t.; CO; 3600 m R. columbianum Cuatrec.; CO; 3100-4200 m R. ecuadorense Jancz.; EC; 3300-4200 m R. hirtum Willd. ex Roem. & Schu1t.; CO, EC; 3350-4400 m R. lehmannii Jancz.; EC; 3500-4350 m R. leptostachyum Benth.; CR?, CO; 3100-4200 m R. peruviana Jancz.; PE; 3400-4050 m R. weberbaueri Jancz.; PE; 3400-3800 m R. weddeliana Jancz. (syn.: R. parvijlorum Wedd.); EC; 3950 m
HALORAGACEAE Gunnera L. Ref.: Mora 0., 1984. G. cf. annae Schind1er; PE; 3300-3450 m G. bogotana L.E.Mora; CO; 2700-3000 m G. caucana L.E.Mora; CO; 2900-3500 m G. garciae-barrigae L.E.Mora; CO; 3700-3900 m G. magellanica Lam.; CO, EC, PE; 3200-4300 m G. pilosa Kunth; CO, EC; 3600-3700 m G. scabra Ruiz & Pav.; CO; 3500-3600 m G. schultesii L.E.Mora; CO; 2800-3400 m G. tacueyana L.E.Mora; CO; 3500 m G. talamancana H. Weber & L.E.Mora; CR; 3100-3400 m G. tamanensis L.E.Mora; CO; 3700 m Myriophyllum L. Ref.: Orchard, 1981. M. aquaticum (Ve!!.) Verde.; CO; 3350 m M. quítense Kunth (syn.: M. elatinoides Gaudich.); CO, VE, EC, PE; 3450-4100 m
HYDROCHARITACEAE Ref.: Cook, 1985.
Elodea Michx. E. matthewsii (Planch.) H.St.John; EC; 3950 m E. potamogeton (Bertero) Espín.; EC; 3900-4100 m
HYDROPHYLLACEAE Phacelia Juss. P. secunda J.F.Gmel.; PE; 3500-4000 m
HYPOXIDACEAE Hypoxis L. H. decumbens L.; CO?, VE, PE; 3000-4000 m H. humilis Kunth; CO; 3250-3350 m
IRIDACEAE Orthrosanthus Sweet O. acorifolius (Kunth) Ravenna; VE; 3400-3700 m O. chimboracensis (Kunth) Baker var. chimboracensis; CR, CO, VE, EC, PE; 3000-4900 m O. monadelphus Ravenna; PA; 3475 m O. occissapungens (Ruiz ex K1att) Die1s; PE; 3200-3900 m Sisyrinchium L. S. alatum Hook.; CO; 2900-3700 m S. brevipes Baker; EC, PE; 4060-4300 m S. chilensis Hook.; EC; 2900-3600 m S. convolutum Nocca; CO, PE; 3200-3900 m S. jamesonii Baker; CO, VE, EC; 3300-4300 m S. micranthum Cav. (syn.: S. iridifolium Kunth); CO, VE, EC; 3000-3600 m S. pusillum Kunth; CO, EC; 3200-4300 m S. subalpinum Henrich & Go1db1att; CR; 1600-3500 m S. tinctorum Kunth (syn.: S. bogotense Kunth); CO, VE, EC; 3300-4100 m S. trinerve Baker; CR, CO, EC, PE; 3000-4300 m S. unispatheum K1att; CO; 3000-3200 m
JUNCACEAE Ref.: Ba1slev, 1996.
Distichia Nees & Meyen D. acicularis Ba1s1ev & Lregaa¡;d; EC; 3200-4200 m D. muscoides Nees & Meyen [syn.: D. tolimensis (Decne.) Buchenau]; CO, EC, PE; 3600-4600 m Juncus L. J. arcticus Willd. var. andicola (Hook.) Balslev; CO, EC, PE; 2700-4200 m J. arequipensis Ba1s1ev; CO; 2700 m J. breviculmis Ba1s1ev; CO, VE; 3500-4100 m J. bufonius L.; CR, CO, VE, EC, PE; 2800-4000 m [Note: Introduced from Europe.] J. capillaceus Lam.; CO, VE, EC; 3450-3500 m J. cyperoides Laharpe; CO, EC, PE; 2800-4000 m J. echinocephalus Ba1s1ev; CO, VE, EC; 3100-4200 m J. ecuadoriensis Ba1slev; CO, VE, EC; 3350-4000 m J. effusus L.; CO, EC; 2700-4000 m J. imbricatus Laharpe; EC; 2500-3600 m J. liebmannii J.F.Macbr. var. quitensis (Buchenau) Bals1ev; CO, EC; 2800-3500 m J. microcephalus Kunth; CO, VE, EC; 1400-3400 m J. pallescens Lam. (syn.: J. dombeyanus J.Gay ex Laharpe); EC, PE; 1900-3600 m J. ramboi Barros subsp. colombianus Ba1s1ev; CO; 26503300m J. stipulatus Nees & Meyen; CO, EC; 2900-4500 m
112
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
J. flimÍds Willd. var. platycaulos (Kunth) Buchenau (syn.: J. andreanus Weath.); CO, EC; 2600-3900 m Luzula DC. L. den ticulata Liebm.; CR, PA; 3100-3500 m L. ecuadoriensis Balslev; EC, PE; 3300-3900 m L. gigantea Desv.; CR?, CO, VE, EC, PE; 2700-4500 m L. racemosa Desv. (syn.: L. peruviana Desv.); CR?, CO, VE, EC, PE; 3100-4875 m L. vulcanica Liebm.; CO, EC; 3400-4800 m Rostkovia Desv. R. magellanica (Lam.) Hook.f.; EC; 3350-4200 m
JUNCAGINACEAE Lilaea Bonpl. L. scilloides (Poir.) Hauman (syn.: L. subulata Humb. & Bonpl.); CO, EC, PE; 3700-4300 m
[VOL. 84
S. styphelus Epling; PE; 3450-4000 m S. tolimensis Kunth; CO; 3400.-3500 m S. xanthophylla Epling & Játiva; PE; 3900 m Satureja L. (syn.: Gardoquia Ruiz & Pav.; Micromeria Benth.) Ref.: Epling & Játiva, 1964. S. caerulescens (Benth.) Epling (syn.: S. lindeniana Briq.); CO; 3300-4800 m S. discolor (Kunth) Briq.; CO; 3050-3100 m S. grisea Epling; CO; 3400-3550 m S.jamesonii (Benth.) Briq.; EC; 3000-4000 m S. nubigena (Kunth) Briq. (syn.: Micromeria nubigena Kunth); CO, VE, EC, PE; 3500-4500 m; includes var. glabrescens Benth. and var. nubigena S. cf. revoluta{Ruiz & Pav.) Briq.; PE; 3600 m s.' sericea (C.Presl ex Benth.) Briq.; PE; 3300-3800 m S. tenella Epling; EC; 3600-4300 m s.·cf. weberbaueri Mansf. (syn.: S. loeseneriana Mansf., S. lopezii Epling); PE; 3400-3500 m
LAMIACEAE Gardoquia Ruiz & Pav. (see Satureja) Lepechinia Willd. L. bullata (Kunth) Epling; CO, VE; 2300-3300 m L. conferta (Benth.) Epling; CO; 3600-3900 m L. salviifolia (Kunth) Epling; CO; 3100-3300 m L. schiedeana (Schltdl.) Vatke [syn.: L. alpina (Oerst.) Stand!., L. hirsuta Epling, L. procumbens Benth.]; CR, CO; 3000-3750 m Micromeria Benth. (see Satureja) Minthostachys (Benth.) Spach M. mollis Griseb.; CO, EC, PE; 3250-3450 m Prunella L. P. vulgaris L.; EC; 2500-4000 m [Note: Introduced from Europe.] Salvia L. Ref.: Wood & Harley, 1989. S. amethystina Sm. subsp. amethystina; CO; 3600 m S. carnea Kunth var. carnea (syn.: S. killipiana Epling); CO, VE, EC; 3300-4100 m S. cocuyana Fern.Alonso; CO; 3000-3750 m S. corrugata Vah!; EC, PE; 2900-3800 m S. cuatrecasana Epling; CO; 3050 m S. hirtella V ah!; PE; 3400-3500 m S. lanicaulis Epling & Játiva; PE; 3500-4000 m S. cf. lobbii Epling; PE; 3400-3550 m S. melaleuca Epling; CO; 3000-3400 m; includes subsp. melaleuca and subsp. totensis J.R.I.Wood & Harley S. nubigena J.R.I.Wood & Harley; CO; 3500-3800 m ' S. palifolia Kunth; CO; 2600-3600 m S. ¡-ubescéns Kunth subsp. truxillensis (Briq.) J.R.I.Wood & Harley; VE; 3200-3400 m
Stachys L. S. bogotensis Kunth; CO; 3100-3 700 m S. elliptica Kunth; CO, VE, EC; 3500-4000 m S. eriantha Benth.; CO, VE, EC; 3000-4200 m S. lamioides Benth.; CO, EC?; 3450 m S. pusilla (Wedd.) Briq.; CO, EC; 3750 m S. repens M.Martens & Galeotti; CO, EC; 3600-4200 m S. venezuelana Briq.; VE; 3000-3600 m
LAURACEAE Perseo Mili. [Note: A forest genus, but rarely found in subpáramo forest patches.] P.ferruginea Kunth; CO, EC; 3048-39.00 m P. mutisii Kunth; CO, VE; 2800-3550 m
LEMNACEAE Lemna L. L. minar L.; CO; 3300 m
LENTIBULARIACEAE Pinguicula L. P. calyptrata Kunth; CO, EC; 3100-4100 m P. diversifolia Cuatrec.; CO; 3300-3500 m P. elongata Benj.; CO, VE; 3100-4450 m P. huilensis Cuatrec.; CO; 3400-3500 m P. involuta Ruiz & Pav.; PE; 3400-3650 m Utricularia L. U. gibba L. (syn.: U. obtusa Sw.); CO; 3650-3900 m
LILIACEAE Eccremis Willd. ex Baker (see Excremis)
:¡.
1999]
PARAMOS
Echeandia Ortega E. ciliata (Kunth) Cruden; CO, VE; 2800-3550 m Excremis Willd. ex Schult.f. E. coarctata (Ruiz & Pav.) Baker; CO; 3250 m Jsidrogalvia Ruiz & Pav. (syn.: Tofie/dia Huds.)
113
Notanthera (DC.) G.Don (see Tristerix longebracteatus) Tristerix Mart. T.longebracteatus (Desr.) Barlow & Wiens [syn.: Notanthera longebracteata (Desr.) G.Don; Phrygilanthus /ongebracteatus (Desr.) J.F.Macbr.]; CO, EC, PE; 2900-4400 m T. secundus (Benth.) Kuijt; CO; 2800-3450 m
Ref.: Cruden, 1991. J. fa/cata Ruiz & Pav. [syn.: Tofie/diafalcata (Ruiz & Pav.) Pers., T. flexuosa Willd., T.frigida Kunth]; CO, EC, PE;
2300-3450 m J. robustior (Steyerm.) Cruden (syn.: Tofieldia sessi/ijlora Hook. var. robustior Steyerm.); VE; 2800-3700 m l. sessilijlora (Hook.) Cruden [syn.: Tofieldia sessi/ijlora Hook.,' T. moritziana (Klotzsch ex Baker) Schulze]; CO,
VE; 2100-4200 m
LYTHRACEAE Cuphaea P.Browne C. ciliata Ruiz & Pav.; CO, PE; 2700-3200 m C. serpyllifo/ia Kunth; CO; 3100 m
MALVACEAE
c.
Acaulimalva Krapov. Tofieldia Huds. (see Jsidrogalvia)
LOASACEAE [Note: The family is not typical to open pรกramo, but is infrequently encountered growing next to large rocks or in the protection of taller vegetation. It is more common to disturbed subpรกramo.] Ref.: Weigend, 1996. Caiophora C.Presl (1831) (not Cajophora Endl., 1839) C. sepiaria (G.Don) J.F.Macbr.; PEยก 4300 m [Note: Weigend
(pers. comm.) believes that thls name is dubious and probably has to be rejected as ~~ synonym. The proper name for this taxon is still in doupt, but may lie near C. carduifolia C.Piesi or C. tenuis Kl(llip.] Loasa Adans. \ L. ranunculifolia Kunth vel aff.; PE; 3\50-4300 m [Note: W,eigend (pers. comm.) believes tha~ollections identified as this taxon in N Peru represen\ a new species and that there are another 2-3 new species yet to be nam~d from this area. Furthermore, he states that L. lindeniana Urb. & Gilg (from VE), L. argemonoides Juss. (from CO), L. karsteniana Urb. & Gilg (from CO), and L. perijensis Weigend (from CO) may also occur occasionally in sheltered subpรกramo si tes.]
LORANTHACEAE [Note: See also Eremolepidaceae and Viscaceae fortaxa formerly placed in Loranthaceae sensu lato.] Ref.: Kuijt, 1986. Aetanthus (Eichler) Engl. A. colombianus A.C.Sm.; CO, VE; 2400-4000 m A. dichotomus (Ruiz & Pav.) Kuijt; CO; 2400-3400 m A. mutisii (Kunth) Engl. [syn.: Psittacanthus holtonii Eichler; A. holtonii (Eichler) Engl. & Prantl]; CO; ? m Gaiadendron G.Don G. punctatum (Ruiz & Pav.) G.Don [syn.: G. lanceolatum (Ruiz & Pav.) Baehni ex J.F.Macbr., G. tagua (Kunth)
G.Don]; CR, PA, CO, VE, EC, PE; 3000-4000 m
Ref.: S. R. Hill, 1982; Krapovickas, 1974. A. acaulis (Dombey ex Cav.) Krapov.; VE; 3700-4100 m A. alismatifolia (K.Schum. & Hieron.) Krapov. (syn.: Malvastrum alismatifolium K.Schum. & Hieron.); PE;
3000-3700 m A. crenata (A.W.Hill) Krapov. (syn.: Malvastrum crenatum
A.W.Hill); PE; 3800-4100 m A. eng/eriana (Ulbr.) Krapov. (syn.: Ma/vastrum eng/erianum Ulbr.); PE; 3200-4100 m A. parnassiaefolia (Hook.) Krapov. [syn.: Malvastrum parnassiaefolium (Hook.) A.Gray]; EC, PE; 3300-4100 m A. purdiaei (A.Gray) Krapov. (syn.: Ma/vastrum purdiaei A.Gray, M. meridae A.W.Hill); CO, VE; 3100-4100 m A. purpurea (A.W.Hill) Krapov. (syn.: Malvastrum purpureum A.W.Hill); CO, VE; 3600-4500 m A. rauhii (Hochr.) Krapov.; PE; 3500-4800 m A. stuebelii (Hieron.) Krapov. (syn.: Malvastrum stuebelii
Hieron.); PE; 3100-3500 m A. sulphurea Krapov.; PE; 3200-3500 m Ma/vastrum A.Gray (see Acaulimalva) Nototriche Turcz.
Ref.: A. W. Hill, 1909. N. artemisioides A.W.Hill; PE; 3900-4000 m N. chimborazoensis Hochr.; EC; 4150 m N. ecuadoriensis Fryxell; EC; 3950-4400 m N. jamesonii A.W.Hill; EC; 4000-4800 m N. cf. longissima A.W.Hill; PE; 4000 m N. lopezii Krapov.; PE; 3800-4000 m N. phyl/anthos (Cav.) A.W.Hill [syn.: N. pichinchensis (Humb. & Bonpl.) A.W.Hill]; EC; 4000-4900 m
MELASTOMATACEAE Ref.: Wurdack, 1973, 1978, 1980. Axinaea Ruiz & Pav. A. macrophylla (Naudin) Triana [syn.: A. affinis (Naudin)
Cogn.]; CO, VE, EC, PE; 3000-3500 m A. merianiae (DC.) Triana [syn.: A. /epidota (Benth.)
Triana]; EC; 2600-3400 m
114
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
Brachyotum (DC.) Triana Ref.: Wurdack, 1953. B. alpinum Cogn.; EC; 3200-4500 m B. andreanum Cogn.; EC, PE; 2700-3400 m B. angustifolium Wurdack; PE; 3300-3500 m B. barbeyanum Cogn.; PE; 2800-3450 m B. campii Wurdack; EC; 3350-3500 m B. cern'uum (Bonpl.) Triana; CO; 3200-3700 m B. cogniauxii Wurdack; PE; 2900-3750 m B. confertum (Bonpl.) Triana; EC; 3100-3500 m B.fictum Wurdack; EC; 3000-3600 m B.figueroae J.F.Macbr.; PE; 3600 m B.fraternum Wurdack; EC; 3400-3500 m B. harlingii Wurdack; EC; 3300-3400 m B. jamesonii Triana; EC; 3000-4000 m B. ledifolium (Desr.) Triana; CO, EC; 2600-4200 m B. lindenii Cogn.; CO, EC; 3200-4200 m B. /ongisepa/um Wurdack; PE; 3300-4000 m B. lymphatum Wurdack; CO; 3300-3800 m B. naudiiJii Triana; PE; 3100-3550 m B. raduĂa Triana; PE; 3000-3700 m B. rostratum (Naudin) Triana (syn.: B. seorsum Wurdack); EC, PE; 3250-4100 m B. rotundifoiium Cogn.; EC; 2840-3800 m B. strigosum (L. f.) Triana; CO; 3050-3900 m Bucquetia DC. B. glutinosa (L.f.) DC.; CO; 2800-3600 m B. vernicosa G1eason; CO, VE; 2900-3000 m Castrate/la Naudin C. pilose/loides (Bonpl.) Naudin; CO, VE; 3050-4200 m C. rosea G1eason; CO, VE; 3100-3700 m Chaeto/epis (DC.) Miq. C. alpina Naudin; CO, VE; 3350-4100 m C. cufodontisii Standl.; CR; 2000-3820 m C. lindeniana (Naudin) Triana [syn.: C. a/pestris (H.Karst.) Triana]; CO, VE; 3200-4900 m C. loricare/la Triana; CO; ? m C. microphy/la (Bonpl.) Miq.; CO, VE; 2700-3700 m C. perijensis Wurdack; CO, VE; 3100-3600 m C. santamartensis Wurdack; CO; 3050-3100 m C. thymifolia Triana; CO; 2800-3000 m Meriania Sw. M tetragona (Cogn.) Wurdack; EC; 2800-3400 m [Note: A marginally subpĂĄramo species at 3400 m.] Miconia Ruiz & Pav. M arbutifolia Naudin; VE; 2800-3300 m M aspergillaris (Bonpl.) Naudin; EC, PE; 3000-3700 m M avia Wurdack; VE; 2900 m M. barclayana Wurdack; EC; 3400-3900 M. biappendiculata (Naudin) L.Uribe; CO; 3200-3400 m M bu/lata (Turcz.) Triana; EC, PE; 3550-3650 m M. buxifolia Naudin; CO, VE; 3300-4100 m M. castillensis Wurdack; EC; 3300-3500 m
m
[VOL. 84
M. cataractae Triana; CO, VE; 2400-2850 m M. chionophila Naudin; CR, PA, CO, VE, EC, PE; 35004300 m M. chlorocarpa Cogn.; CO, EC; 2800-3700 m M. cladonia Gleason; EC; 3200-3350 m M cleefii L.Uribe; CO; 3450-3750 m M cundinamarcensis Wurdack; CO; 3000-3200 m M elaeoides Naudin; CO, VE; 2900-3600 m M. elvirae Wurdack; VE; 2900 m M gleasoniana Wurdack; CO; 3200-3500 m M. griffisii J.F.Macbr.; PE; 3550 m M. harlingii Wurdack; CO, EC; 3400-3650 m M hexamera Wurdack; EC; 3100-3450 m M.jahnii Pittier; CO, VE; 2900-3300 m M. jentaculorum Wurdack; CO; 3100-3200 m M /atifolia (D.Don) Naudin (syn.: M. andina Naudin, M. epiphytica Cogn.); CO, VE, EC, PE; 3300-4050 m M /edifolia (DC.) Naudin; EC; 2850-3600 m M ligustrina (Sm.) Triana var. setulinodis Wurdack; CO, EC; 3300-3700 m M limitaris Wurdack; CO, VE; 3000-3100 m M. media (D.Don) Naudin subsp. borealis Wurdack; PE; 2800-3500 m M. mesmeana G1eason; CO, VE; 3100-3600 m; ineludes subsp.jabonensis Wurdack and subsp. mesmeana M. myrtil/ifolia Naudin; CO, VE; 3000-3200 m M nitidissima Cogn.; VE;.2720-2900 m M. nodosa Cogn.; CO; 3000-3250 m M. orcheotoma Naudin; CO; 3000-3600 m M. oreogena Wurdack; CO; 3250-3350 m M. paZ/ida Gleason; CO; 3220-3900 m M. paludigena Wurdack; PE; 3500-3750 m M. parvifolia Cogn.; CO; 3250-3500 m M. pernettifolia Triana; EC; 3400-3500 m M polyneura Triana; CO; 3300-3350 m M puracensis Wurdack; CO; 3500-3700 m M. radula Cogn.; PE; 2900 m M. rigens Naudin; CO; 3200-3300 m M rotundifolia (D.Don) Naudin; EC, PE; 3400-3750 m M. salicifolia (Bonpl. ex Naudin) Naudin; CO, EC, PE; 2950-4150 m M. schnellii Wurdack; CR; 3050-3200 m M. spinulidentata Cogn. & Gleason ex G1eason; CO; 38004000m M squamulosa (Sm.) Triana; CO; 2800-3000 m M stipularis Naudin; CO; 2400-3400 m M. summa Cuatrec.; CO; 3200-3700 m M tephrodes Wurdack; EC; 3350-3450 m M. tinifolia Naudin; CO, VE, EC; 2500-4100 m; inc1udes var. parviflora Cogn. and var. tinifolia M. tricaudata Wurdack; CO; 3250-3350 m M. ulmarioides Naudin; VE; 2800-3300 m M. vaccinioides (Bonpl.) Naudin; PE; 3400 m M. verrucosa Cogn.; CO; 3200 m Monochaetum (DC.) Naudin M. amistadense Almeda; CR; 2400-3300 m M.bonplandii (Kunth) Naudin; CO, VE; 2300-3800 m
1999]
M. M. M. M. M. M. M. M. M. M.
PARAMOS
coronatum Gleason; CO; 2800-3100 m discolor H.Karst.; VE; 2900-3100 m glanduliferum Triana; CO; 2900-3100 m gleasonianuin Wurdack; VE; 2700-3300 m mariae Wurdack; VE; 3100-3200 m myrtoideum (Bonpl.) Naudin; CO; 2900-3650 m paucijlorum Triana; CO, EC; 3200-3300 m rodriguezii W urdack; VE; 2100-2900 m stellulatum Naudin; CO; 3000-3100 m uberrimum Sandwith; CO; 3000 m
Tibouchina Aubl. T andreana Cogn.; CO; 3200-3650 m T. grossa (L.f.) Cogn. [syn.: Melas/ama grossa L.f.; T. reticulata (Bonpl.) Cogn.]; CO, VE, EC; 3000-4100 m T mollis (Bonpl.) Cogn.; CO; 2300-3700 m; includes var. glandltlifera Wurdack and var. mollis T. stricta Wurdack; CO; 3300-3800 m
MYRICACEAE Myrica L. M..funckii Chev.; CO, VE; 3200-3300 m M. parvifolia Benth.; CO, VE, EC; 3000-3700 m M. pubescens Humb .. & Bonpl ex Willd.; CO, EC; 18003600m
MYRSINACEAE Conomorpha DC. (see Cybianthus) Cybianthus Mart. (syn.: Conomorpha DC., Grammadenia Benth.) Ref.: Pipoly, 1987. C. baruana (see Comarostaphylos arbutoides in the Ericaceae) C. iteoides (Benth.) G.Agostini subsp. nevadensis (Mez) Pipoly; VE; 2950 m C. marginatus (Benth.) Pipoly (syn.: Grammadenia alpina Mez); CO, VE, EC; 2800-4000 m C. perseoides (Mez) G.Agostini; CO; 3300-3400 m C. stapfii (Mez) G.Agostini; CO; 2700-3500 m Geissanthus Hook.f. G. andinus Mez; CO, VE; 2800-3700 m G. quindiensis Mez; CO, EC; 2800-3600 m Grammadenia Benth. (see Cybianthus) Myrsine L. (syn.: Rapanea Aubl.) Ref.: Pipoly, 1992. M. andina (Mez) Pipoly; CO, EC; 2800-4000 m M. cm'iacea (Sw.) R.Br. ex Roem. & Schult. (syn.: M.jelskii , Zahlbr.); CO; 2800-3200 m M. dependens (Ruiz & Pav.) A.Spreng. [syn.: Rapanea pittieri Mez; Myrsine pittieri (Mez) Lundell]; CR, PA, CO, VE, EC, PE; 2750-4100 m M. umbellata Mart.; EC; 3200-3300 m
115
Rapanea Aubl. (see Myrsine)
MYRTACEAE Eugenia L. E. triquetra O.Berg (syn.: E. weberbaueri Diels); CO, VE, PE; 3200-3350 m; disturbed areas Myrcianthes O.Berg M. leucoxyla (Ortega) McVaugh; CO; 2700-3100 m M. rhopaloides (Kunth) McVaugh [syn.: Eugenia rhopaloides (Kunth) DC.]; VE; 2500-3400 m Myrteola O.Berg Ref.: Landrum, 1988. M. acerosa (O.Berg) Burret; PE; 2900-3700 m M. hummularia (Poir.) O.Berg [syn.: M. oxycoccoides (Benth.) O.Berg, M. vaccinioides (Kunth) O.Berg]; CO, VE, EC, PE; 3300-4200 m M. phylicoides (Benth.) Landrum var. glabrata (O.Berg) Landrum [syn.: M. microphylla (Humb. & Bonpl.) O.Berg var. glabra/a O.Berg]; PE; 2800-38\:J~ Ugni Turcz. U. myricoides (Kunth) O.Berg [syn.: U. angustifolia Burret, U. montana (Benth.) O.Berg, U. oerstedii (O.Berg) Hemsl., U. warscewiczii O.Berg]; CR, PA, CO, VE; 3100-3700m
NYCTAGINACEAE Colignonia Endl. Ref.: Bohlin, 1988. C. ovalifolia Heimerl; CO; 3300-3600 m C. parvijlora (Kunth) Choisy subsp. biumbellata (Ball) Bohlin; PE; 3300-4000 m
ONAGRACEAE Epilobium L. Ref.: Solomon, 1982. E. denticulatum Ruiz & Pav. (syn.: E. aequinoctiale Sam., E. meridense Hausskn.); CR, CO, VE, EC, PE; 32004650 m Fuchsia L. [Note: A forest genus, but frequently getting into disturbed weedy situations in subpรกramo.] Ref.: Berry, 1985. F. ampliata Benth.; EC; 2900-3475 m F. ayavacensis Kunth; EC; 3500 m F. canPscens Benth.; CO; 3260 m F. cf. caucana P.E.Berry; CO; 2950-3300 m F. colombiana Munz; CO; 3500-3600 m F. con.fertifolia Fielding & Gardner; PE; 3100-3300 m F. corollata Benth.; CO, EC; 2900-3450 m F. dependens Hook.; CO; 2900-3400 m F..fontinalis J.F.Macbr.; PE; 2900-3400 m
116
MEMOIRS OF THE NEW YORK BOTAN! CAL GARDEN
' gehrigeri Munz; CO, VE; 2200-3400 m /oxensis Kunth; EC; 3250-3800 m magdalenae Munz; CO; 3000-3350 m mathewsii J.F.Macbr.; PE; 280-3400 m membranacea Hemsl.; VE; 2800-3400 m cf. osgoodii J.F.Macbr.; PE;? m petiolaris Kunth; CO; 2750-4100 m sanmartina P.E.Berry; PE; 3100-3650 m sessilifolia Benth.; CO, EC; 2500-3500 m splendens Zucc.; CR; 3000-3800 m F. summa P.E.Berry; EC; 3100-3450 m F. venusta Kunth (syn.: F. killipii I.M.Johnst.); CO; 28004000 m F. vu!canica André; CO, EC; 3100-3500 m F. F. F. F. F. F. F. F. F. F.
Oenolhera L. O. epilobiifolia Kunth (syn.: O. cuprea Schlecht.); CR, CO, VE, EC; 2900-3950 m; inc!uding subsp. cuprea (Schlecht.)P.H.Raven & J.Parn. and subsp. epilobiifolia O. mullicaulis Ruiz & Pav. (syn.: O. larquensis Kunth); CR, VE, EC, PE; 2900-3800 m O. seifrizii Munz; CO, VE; 2800-4500 m O. versicolor Lehm.; PE; 3300-4000 m
ORCHIDACEAE Aa Rchb.f. [Note: Dodson (pers. comm.) lists Aa riobambae Schltr. from EC at 3 800 m, but 1 have not seen any voucher nor do l have !abe] information about exact localities.] A. argyrolepis Rchb.f.; CO, EC; 4100-4500 m A. colombiana Schltr. [syn.: Allensleinia colombiana (Schltr.) Garay]; CO, EC; 3800-4300 m A. denticulata Schltr.; CO, EC; 2900-3900 m A. hartwegii Garay; VE, EC; 3800-4100 m A. leucantha (Rchb.f.) Schltr.; CO, VE, EC; 3500-4200 m A. maderoi Schltr. (syn.: A/tensteiniafragosa Lojtnant); CO, EC; 3300-4300 m A. paleacea (Kunth) Rchb.f. [syn.: Altensleinia pa/aceae (Kunth) Kunth]; CR, CO, VE, EC, PE; 3000-3900 m A. rhynchocarpa Schltr.; CO; 4400 m A. rostrata (Rchb.f.) Schltr. [syn.: Myrosmodes rostratum (Rchb.f.) Garay]; CO, VE; 3800-4400 m Altensteinia Kunth [Note: Many older Altensteinia names have now been transferred to Aa or Myrosmoides.] A.fimbriata Kunth; CO, VE, PE; 2600-3500 m A. lemantha H.B.K.; CO; 3400-3500 m A. virescens Lindl.; EC; 2800-3550 m Barbosella Schltr. B. cucullata (Lindl.) Schltr.; CO; 3500 m Brachionidium Lindl. [Notes: The species of Brachionidium are often found on road embankments in páramo. Luteyn el al. 12837 (Brachionidium sp., COL, NY) is determined only
[VOL. 84
to genus by E. Christenson, but would be the first record ofthis genus noted from Colombia.] B. tetrapetalum (F.Lehm. & Kraenzl.) Schltr.; EC; 37003850m B. tuberculatum Lindl.; EC; 3750 m Cranichis Sw. [Note: A forest genus, but rarely found in subpáramo., Dodson (pers. comm.) lists Jorgl!nsen el al. 92261 (C. sp., MO) from EC at 3460 m as occurring in páramo.] C. cf. cucullata Schltr.; CO; 3200 m C. diphylla Sw.; CO; 3150 m Elleanlhus C.Presl E. auranliacus (Lindl.) Rchb.f. [syn.:? E.jlavescens (Lindl.) Rchb.f.]; CO, VE, EC; 2500-3700 m E. ensatus (Lindl.) Rchb.f.; CO; 2800-4000 m E. gracilis (Rchb.f.) Rchb.f.; EC; 3400-3900 m E. scopulus Schltr.; EC; 2900-3700 m E. ventricosus Schltr.; EC; 3300--4100 m E. vinosus Schltr.; CO; 3300-3500 m E. virgatus (Rchb.f.) C.Schweinf.; EC; 4100 m E. wercklei Schltr.; PA; 3100-3300 m Epidendrum L. E. acuminatum Ruiz & Pav.; PE; 3100-3250 m. E. aquaticoides C.Schweinf.; EC; 3000-3420 m E. brevivenium Lindl.; EC; 2800-3400 m E. chioneum Lindl.; CO, VE; 3100-3500 m E. chortophyllum Schltr.; EC; 3400-3800 m E. cuniculatum Schltr.; EC; 3800 m E. dermatanthum Kraenzl.; EC; 4100 m E. elleanthoides Schltr.; CO, EC; 3000-3750 m E. erosum Ames & C.Schweinf.; CO; 3000-3600 m E. fimbriatum Kunth; CO, VE, EC, PE; 2800-4000 m E.frigidum Linden ex Lindl.; CO, VE, EC; 3000-4450 m E.frutex Rchb.f.; CO, VE; 2950-4000 m E. gaslropodium Rchb.f.; EC; 2000-3500 m [Note: A species found along road embankments in páramo.] E. globiflorum F.Lehm. & Kraenzl.; VE, EC; 2750-3500 m E. gramineum Lindl.; PE; 3300-3500 m E. inornatum Schltr.; EC; 3650-3750 m [Note: A species found along road embankments in páramo.] E. loxense F.Lehm. & Kraenzl.; EC; 2900-3150 m E. macrostachyum Lindl.; EC; 2000-3700 m E. microdendron Rchb.f.; CR; 3100-3400 m E. moronense Dodson & Hágsater; EC; 3500 m E. paucifolium Schltr.; CR; 3100-3400 m E.pichinchae Schltr.; EC; 2500-3800 m E. rhombochilum L.O.Williams; EC; 3300-3850 m E. scabrum Ruiz & Pav.; EC; 1900-3700 m E. serpens Lindl.; CO, EC?; 3000-3530 m E. torquatum Lindl.; CO; 2800-3400 m E. vesicicau/e L.O.Williams; EC; 3600-3800 m Gomphichis Lindl. G. bogolensis Renz; CO, VE; 3000-3700 m
'/
1
1999]
PARAMOS
G. brachystachys Schltr.; CO; 3150 m G. caucana Schltr.; CO, VE, EC; 3000-3500 m G. crassilabia Garay; EC; 3450-3500 m G. cundinamarcae Renz; CO; 3000-3500 m G.foliosa Ames; CO, VE; 3000-3500 m G. goodyeroides Lindl.; EC; 2900-3650 m G. hetaerioides Schltr.; EC; 1800-4000 m G. macbridei C.Schweinf.; EC; 3600-3700 m G. montana L.O.Williams; VE; 3100-3300 m G. traceyae Rolfe; CO, VE, EC; 2900-4000 m G. valida Rchb.f.; EC; 2600-4000 m G. viscosa (Rchb.f.) Schltr.; VE, PE; 2800-3700 m Habenaria Willd. H. gollmeri Schltr.; CO, EC; 2500-3700 m Lepanthes Sw. [Notes: Severa! species of Lepanthes-e.g., L. amplisepala Luer & R.Escobar, L. deliciasensis Luer & R. Escobar, L. guanacasensis Luer & R.Escobar, and L. os iris Luer & R.Escobar-have been reported (in lit.) as occurring in the scrub forestofvarious páramos in central Colombia at 3150-3380 m; however, I have not seen voucher specimens, nor is exact !abe! information available. Dodson (pers. comm.) also lists L. biloba Lindl., L. chimaera Luer & R.Escobar, L. decurva Luer & Hirtz, L. de/phax Luer, L. frigida Luer & Hirtz, L. golbasto Luer & Hirtz, L. il/ex Luer, L. mastix Luer, L. metaxy Luer & Hirtz, L. monitor Luer, L. monr.¡ptera Lindl., L. muscula Luer & R. Escobar, L. otostalyx Rchb.f., and L. oxypetala Luer & Hirtz from EC at 3400-3600(-3700) m, but I have not seen any voucher material nor do I have exact !abe! information; these are probably epiphytes in high montane forest, not in páramo.] L. aberrans Schltdl.; EC; 3500-3850 m L. altico/a Schweinf.; EC; 3600 m L. deliqua Luer; EC; 3600 m L. dolichopus Schltr.; CO, EC; 3450-3600 m L. gargantua Rchb.f.; CO; 3200-3400 m L. mesochlora Rchb.f.; CO; 3000-3200 m L. mucronata Lindl.; EC; 3450-3600 m L. rhombipetala Schltr.; EC; 3450 m L. tachirensis Foldats; CO; 3650-4290 m L. trachysepala Schltr.; EC; 3800-3850 m L. tricuspis Schltr.; CO; 3300-3350 m Ma/axis Sol. ex Sw. [Notes: A forest genus, but also found in shrub páramo. Dodson (pers. comm.) does list M. andico/a (Ridl.) Kuntze, M. caracasana (Klotzsch ex Ridl.) Kuntze, M. excavata (Lindl.) Kuntze, and M. hoppii (Schltr.) Lojtnant from EC at 3400-3600 m and M. sodiroi (Schltr.) Dodson from 3850 m, but I have not seen any voucher material not do I have exact !abe! information; these are proba! y epiphytes in high montane forest, not in páramo.] M. soulei L.O.Williams; CR, VE; 1000-3500 m
117
Masdevallia Ruiz & Pav. [Notes: A forest genus, but sometimes found on rocks or as a terrestrial amongst mosses in páramo. Dodson (pers. comm.) does list M. ajjinis Lindl., M. anachaeta Rchb.f., M. pardina Rchb.f., and M. uncifera Rchb.f. from EC at 3400-3600 m, but I have not seen any voucher material nor do I have exact !abe! information; these are probably epiphytes in high montane forest, notin páramo.] M. amabilis Rchb.f. & Warsw.; PE; 2900-3750 m M. corderoana F.Lehm. & Kraenzl.; CO, EC; 3100-3400 m M. coriacea Lindl.; CO, EC; 3100-3900 m; includes var. bonplandii (Rchb.f.) Luer and var. coriacea [Note: Often growing on rocks.] M. ignea Rchb.f.; CO; 3550-3800 m M. semiteres Luer & R.Escobar; PE; 3100-3800 m [Note: Growing on rocks.] Maxil/aria Ruiz & Pav. [Note: Dodson (pers. comm.) does list M. alticola C.Schweinf. from EC at 3500 m, but I have not seen any voucher material nor do I have exact !abe! information; this is probably an epiphyte in high montane forest, not in páramo.] M. aurea (Poepp. & Endl.) L.O.Williams; CO, PE; 28003500 m M. /epidota Lindl.; EC; 4100-4500 m M. pleuranthoides (Schltr.) Garay: CO; ? m M. pulla Linden & Rchb.f.; PE; 3100-3250 m Myrosmodes Rchb.f. M. brevis (Schltr.) Garay; EC; 3450-3900 m M. coch/eare Garay; VE, EC; 3200-4400 m M. nubigenum Rchb.f. [syn.: Altensteinia nubigena (Rchb.f.) Rchb.f.]; VE, EC; 3000-4500 m M. paludosa (Rchb.f.) Garay (syn.: Aa pa/udosa Schltr.; Altensteinia paludosa Rchb.f.); CO, VE, PE; 3625-4600 m M. rhynchocarpum (Schltr.) Garay; EC; 3200-4000 m M. ustulatum (Schltr.) Garay; EC; 3000-4100 m Odontoglossum Kunth [Note: Dodson (pers. comm.) does listO. auropurpureum Rchb.f. from EC at 3400-3500 m, but I ha ve not seen any voucher material nor do I have exact labe1 information.] O. angustatum Lindl.; PE; 3100-3500 m O. aureum (Lindl.) Rchb.f.; EC; 3500 m O. cristatum Lindl.; EC; 4100 m O. lindenii Lindl.; CO, VE; 3100-3650 m O. oerstedii Rchb.f. (see Ticog/ossum) O. pardinum Lindl.; CO, EC; 3350-4000 m O. prasinum Linden & Rchb.f.; CO; 3200-3300 m O. mmosissimum Lindl.; CO, EC; 3000-3800 m O. retusum Lindl.; EC; 3000-3150 m O. revolutum Lindl.; CO; 3200-3750 m Oncidium Sw. O. aureum Lindl.; EC; 3400 m O. cucul/atum Lindl.; EC; 3400-3750 m
118
MEMOIRS OF THE NEW YORK BOTAN! CAL GARDEN
Pa~h;phyllum Kunth [Note: Dodson (pers. comm.) does 1ist P. peperomoides Kraenzl. from EC at 3400-3500 m, but I have not seen any voucher material nor do I have exact label information.] P. crystallinum Lindl.; EC; 3200-4000 m P. distichum Kunth; PE; 2750-3600 m P. hartwegii Rchb.f.; EC; 2800-3450 m P. lycopodioides Schltr.; PE; 3600 m P. cf. nubivagun; L.O.Williams; CO; 3800-4100 m P. pastii Rchb.f.; CO, EC; 3400-4200 m P. squarrosum Lindl.; CO; 3300-3750 m Platystele Schltr. [Note: Dodson (pers. comm.) does 1ist P. piscifera (Lindl.) Luer and P. stonyx Luer from EC at 34003600 m, but I have not seen any voucher material nor do I have exact 1abel information.] Pleurothallis R.Br. [Notes: Dodson (pers. comm.) dóes list P. adonis Luer, P. apopsis Luer, P. atacasana Luer, P. avirostra Luer & Hirtz, P. nivalis Luer, and P. patula Schltr. from EC at 3400-3600 m and P. praealta Luer & Hirtz from 3 800 m, but I have not se en any voucher material nor do I ha ve exact !abe! information.] P. acuminata Lindl.; VE; 2200-3200 m P. aurea Lindl.; VE; 3050-3475 m [Note: A species found on limestone outcrops.] P. cassidis Lindl.; CO; 3300-3350 m P. declivis Lindl.; EC; 3600-3800 m P.jamesonii Lindl.; CO; 3300-3350 m P. laevigata Lindl.; EC; 3100-3600 m P. ligulata Lindl.; EC; 1800-3600 m P. litotes Luer; EC; 3400 m [Note: A species found on road embankments in páramo.] P. lloensis Schltr.; EC; 3800 m P. pilifera Lindl.; EC; 3400 m [Note: A species found on road embankments in páramo.] P. ramificans Luer; EC; 3400 m [Note: A species found on road embankments in páramo.] P. secunda Poepp. & Endl.; CO; 3000-3400 m P. simplex Ames & C.Schweinf.; CR; 3100-3400 m P. spiralis (Ruiz & Pav.) Lindl.; EC; 2600-3500 m P. talpinaria Rchb.f.; CO; 3200-3500 m Pterichis Lindl. Ref.: Morales L, 1986. P. colombiana G.A.Morales; CO; 3800 m P. 'galeata Lindl. var. galeata [syn.: P. acuminata Schltr.,? P. barbifrons (Kraenzl.) Schltr.]; CO, EC, PE; 31503850 m P. habenarioides (F.Lehm. & Kraenzl.) Schltr. (syn.: P. costaricensis Ames & C.Schweinf.); CR, CO, EC; 22003900m P. leo L.D.Gómez & Gómez-Laur.; CR; 3000-3500 m P. multiflora (Lindl.) Schltr.; CO, VE, EC; 3000-3850 m P. parvifolia (Lindl.) Schltr.; CO; 3150-3350 m
[VOL. 84
P. pauciflora Schltr.; CO; 3300-3700 m P. triloba (Lindl.) Schltr.; CO, EC; 3200-3700 m
Salpistele Dressler S. pensilis (Schltr.) Luer;· EC; 3350-3450 m Spiranthes Rich. (see Stenorrhynchos) Stelis Sw. [Notes: A forest genus, but rarely found in subpáramo forest patches. Dodson (pers. comm.) does list S. polycldda Lindl. and S. purpurea (Ruiz & Pav.) Willd. from EC at 3500 m, but I have not seen any voucher material nor do I ha ve exact label information.] S. apiculata Lindl.; EC; 3500 m S. dialissa (Lindl.) Rchb.f.; CO; 3250 m S.furfuracea Lehm. & Kraenzl.; CO; 3200 m S. humilis Lindl.; VE; 2000-3200 m [Note: A.sQecies found on limestone outcrops.] ~ S. lanceo/ata (Ruiz & Pav.) Willd.; VE; 2700-3200 m S. muscosa Lindl.; EC; 3600 m S. pusilla Kunth; CO, EC; 2000-3900 m S. sirio/ata Lindl.; VE; 3000-3150 m Stenorrhynchos Rich. ex Spreng. (syn.: Coccineorchis Schltr.; Spiranthes p.p.) S. vaginatum (Kunth) Spreng. [syn.: Spiranthes vagina/a (Kunth) Lindl. ex B.D.Jackson]; CO, VE; 2750-3550 m Telipogon Mutis ex Kunth [Note: Dodson (pers. comm.) does list T. vollesii Dodson & R.Escobar and one other collection (Hirtz & Hirtz 5117, RPSC) from EC at 3400 m, but I have not seen any voucher material nor do I have exact !abe! information.] T nervosus (L.) Druce; CO; 3200 m T semipictus Rchb.f. ex Kraenzl.; PE; 3100-3250 m T storkii Ames & C.Schweinf.; CR; 3100-3400 m T tessellatus Lindl.; PE; 3250 m Ticoglossum Lucas Rodr. ex Halb. T oerstedii (Rchb.f.) Lucas Rodr. ex Halb. (syn.: Odontoglossum oerstedii Rchb.f.); PA; 2300-3450 m Trichosalpinx Luer [Note: Dodson (pers. comm.) does líst T brevispicata C.Schweinf. and T chaemaelepanthes (Rchb.f.) Luer from EC at 3600 m, but I ha ve not seen any voucher material nor do I have exact !abe! information.] T. microcharis (Schltr.) Luer; EC; 3800 m T pusilla (Kunth) Luer [syn.: Pleurothallis pusilla (Kunth) Lindl.]; CO, VE; 3000-3400 m T quitensis (Rchb.f.) Luer; EC; 4000 m
OXALIDACEAE Oxalis L. O. fendleri Lourteig; CO, PE; 3100-3600 m
1999]
PARAMOS
O. filiformis R.Knuth; CR, CO, EC; 3600--4000 m O. /otoides K.unth; CO, EC, PE; 3000-3750 m O. medicaginea R.Knuth; CO, VE, EC; 3100--4000 m O. pennelliana R.K.nuth; CO; 3600-3700 m O~ phaeotricha Die1s; CO; 3500--4200 m O. rufescens Turcz.; EC; 3600--4200 m O. spiralis Ruiz & Pav. ex G.Don; CR, CO, VE; 3100-3700 m; inc1udes subsp.-rlpiralis and subsp. vulcanicola (Donn.Sm.) Lourteig O. tabaconasensis R.Knuth; CO; 3200-3700 m O. teneriensis R.Knuth; EC; 4300 m
119
P. P. P. P. P. P. P. P. P. P.
mandonii C.DC.; CO; 3300-3700 m microphylla Kunth; CO, VE, EC; 2950-3900 m mo!lis Kunth; VE; 3100-3500 m parasítica C.DC.; CO, EC; 3300-3350 m peruviana (Miq.) Dahlst.; CO, VE, PE; 3300-3700 m quadrangifolia (L.) Kunth; CO; 2700-3000 m quadrifolia (L.) Kunth; CR; 3100-3400 m quindioensis Trel. & Yunck.; CO; 3150-3200 m rotundata Kunth; CO; 3300-3790 m sa/igna Kunth (syn.: P. bistortaefolia Trel., P. alagotacta C.DC.); CR, PA, CO, VE; 2800--4000 m P. tequendamana Trel.; CO, VE; 2800--4000 m
PASSIFLORACEAE Ref.: Escobar, 1988; Holm-Nie1sen et al., 1988.
Passijlora L. [Note: A genus found mostly in forest and disturbed are as in subpáramo forest patches.] .f. adulterina L.f.; CO; 3300-3500 m P. bicuspidata (H.Karst.) Mast.; CO; 3000-3200 m P. colombiana L.K.Escobar; CO; 3150-3600 m P. crispo/anata L.Uribe (syn.: P. boyacana Killip); CO; 2600-3100 m P. cuatrecasasii K.illip; CO; 2200-3400 m P. cumbalensis (H.Karst.) Harms; CO, EC; 1800--4100 m; inc1udes var. caucana L.K.Escobar, var. cumbalensis, 1 and var. goudotiana (Triana & P1anch.) L. K. Escobar P.jamesonii (Mast.) Bailey; EC; 2800-3750 m P. cf. /anata (Juss.) Poir.; CO; 2200-3500 m P. toxensis Killip & Cuatrec.; EC; 2200-3400 m P. mathewsii (Mast.) K.illip; EC, PE; 2850-3550 m 1 P. mixta L. f.; CO, EC; 1700-3900 m; includes var. eriantha (Benth.) Killip and var. mixta P. parvifolia (DC.) Harms; PE; 2700-3600 m P. schlimiana Triana & Planch.; CO; 2400-3900 m P. sierra e L.K.Escobar; CO; 3100 m P. trianae Killip; CO; 3000-3300 m P. trinervia (Juss.) Pers.; CO; ca. 3500 m
PHYTOLACCACEAE Phyto/acca L. P. australis Phi!.; VE; 3800--4100 m P. bogotensis Kunth; CO, EC; 3500-3800 m P. rugosa Braun & Bouché; CR, PA, CO; 2000-3500 m P. sanguínea H.Walter; CO; 3000-3600 m
PIPERACEAE Peperomia Ruiz & Pav. P. alibacophylla Trel. & Yunck.; CO; 3300-3400 m P. galioides Kunth (syn.: P. jamesonii Regel., P. fiagelliformis Hook.f. ex Miq.); CR, CO, EC; 3100--4000 m P. hartwegiana Miq. (syn.: P. cinerea Sodiro, P. dolichostachya Sodiro, P. kunthiana C.DC.); CO, VE, EC, PE; 3300--4300 m P. hispidula (Sw.) A.Dietr.; CO, EC; 3200-3500 m P. ioeides Trel. & Yunck.; CO; 3250 m
PiperL. [Note: A forest genus, but rarely in subpáramo forest patches.] P. barbatum Kunth; EC; 2300-3260 m P. nubigenum Kunth var. brevifolium Trel. & Yunck.; CO; 2800-3300 m P. ramosense Yunck.; CO; 3150-3200 m
PLANTAGINACEAE Plan lago L. P. alopecuris Decne.; CO; 3600-3700 m P. australis Lam.; CR, CO, VE, EC, PE; 2800--4200 m; ineludes subsp. ecuadorensis (~ilg.) Rahn, subsp. hirtella (Kunth) Rahn, subsp. orea des (Decnc.) Rahn, and subsp. sodiroana (Pi1g.) Rahn P. caridna Decne.; CO; 2800-3900 m P. linearis Kunth; CO, EC, PE; 2500-4500 m; includes var. agrostophylla (Decne.) Pi1g., var. leptotricha Pilg., and var. linearis P. monticola Decne. subsp. angusta Pi1g.; CO; 3500--4150 m P. nubigena K.unth; CO, EC; 3500--4600 m P. rigida Kunth; CO, VE, EC, PE; 3300--4500 m P. sericea Ruiz & Pav.; CO, VE, EC, PE; 3300--4250 m; includes subsp. argyrophylla (Decne.) Rahn and subsp. perrymondii (Barnd.) Rahn P. tarattothrix Pi1g.; PE; 4000--4100 m P. tubulosa Decne.; CO, EC, PE; 3450--4450 m
POACEAE Ref.: Barkworth, 1990; Hitchcock, 1927; Nicora & Rúgo1o de Agrasar, 1987; Poh1, 1977; Tovar, 1993.
Aciachne Benth. Ref.: Lregaard, 1987. A. acicularis Lregaard; CR, CO, VE, EC, PE; 3500--4150 m A.jlagellifera Lregaard; CO, EC; 3850--4400 m A. pulvinata Benth.; CR, VE, EC, PE; 3500--4400 m Aegopogon Humb. & Bonpl. ex Willd. A. cenchroides Humb. & Bonpl. ex Willd.; CO, VE; 25004200m Agropyron Gaertn. (Andean taxa now mostly under Elymus)
120
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
'
Agrostis L. A. araucana Phi!.; CO, EC; 3800-4200 m A. bacillata Hack.; CR; 3050-3800 m A. boyacensis Swallen & García-Barr.; CO, EC; 3700-4500 m A. brevicu/mis Hitchc.; CO;VE, EC, PE; 3300-4650 m A. fascicu/ata (Kunth) Roem. & Schult. [syn.: A. humboldtiana Steud., A. perennans (Walt.) Tuck]; CR, CO, VE, EC, PE; 3000-4000 m ' A. fo/iata Hook.f. (syn.: A. nigritel/a Pilg.); CO, VE, EC, PE; 2800-4800 m A. ge/ida Trin.; Có; 3200 m A. gracilis Pilg.; CO; 4300 m A. haenkeana Hitchc.; CO, VE, EC, PE; 3350-4200 m A.jahnii Luces; VE; 3000-3150 m A.jamesoniana Steud.; EC; 4600 m A. /ehmannii Swallen; CO; 3500 m A. meridensis Luces; VE; 3500-3600 m A. mertensii Trin. (syn.: A. boliviana Mez); EC, PE;? m A. pittieri Hack.; VE; 4000 m A. scabrifolia Swallen; CO; 3100-3200 m A. sodiroana Hack.; EC; 3600-3800 m A. stolonifera L.; EC, VE; 3250 m A. subpatens Hitchc.; CR, VE; 3000-4000 m A. tenuis Sibth.; VE; 3300 m A. tolucensis Kunth (syn.: A. virescens Kunth); CR, PA, CO, VE, EC, PE; 3200-4800 m A. trichodes (Kunth) Roem. & Schult. (syn.: A. bogotensis Hack.); CO, VE, EC, PE; 3100-4000 m A. turrialbae Mez; CR, CO; 2950-3500 m A. venezuelana Mez; VE; 3250-3600 m
[VOL. 84
Brachypodium P.Beauv. B. mexicanum (Roem. & Schult.) Link; CR, CO, VE, EC, PE; 3150-3800 m Eriza L. ( B. minar L.,; CO; 3000-3200 m [Note: Introduced from Europe.] Bromus L. Ref.: Pinto-Escobar, 1981. B. carinatus Hook. & Arn.; CR; 1500-3500 m B. catharticus Vahl (syn.: B. unio/oides Kunth, B. wil/denowii Kunth); CO, EC, PE; 3200-4400 m [Note: This species is also cultivated as forage.] B. co/oratus Steud.; CO; 2750-3300 m 8,. exaltatus Bemh.; CR; 3000-3500 m .B. /anatus Kunth [syn.: B. oliganthus Pilg., B. pitensis Kunth (fide S. Renvoize, pers. comm.)]; CO, VE, EC, PE; 3000-4750 m [Note: This species is native but is sometimes cultivated as forage.] B. segetum Kunth; CO, PE; 3400-3520 m
Ca/amagrostis Adans. (syn.: Deyeuxia Clarion ex P.Beauv.) Ref.: Escalona, 1988; Lregaard, 1998; Tovar, 1960. C. aurea (Munro ex W edd.) Hack. ex Sodiro (syn.: Deyeuxia aurea Munro ex Wedd.); EC; 4200-4800 m C. bogotensis (Pilg.) Pilg. (syn.: Deyeuxia bogotensis Pilg.); CO, VE, EC, PE; 3000-4350 m C. boyacensis Swallen & García-Barr.; CO; 4000-4300 m C. carchiensis Lregaard; EC; 3200-3900 m C. chasei Luces; VE; 3470-4000 m C. c/eefii Escalona; CO; 4100 m A/opecurus L. C. coarctata (Kunth) Steud. (syn.: Deyeuxia coarctata A. aequa/is Sobo!.; CO, VE, EC, PE; 3200-4500 m Kunth, D.fuscata C.Presl); CO, VE, EC; 3650-4700 m C. divergens Swallen; CO; 3100-3200 m Andropogon L. (see Bothriochloa saccharoides) C. ecuadoriensis Lregaard; EC; 3450-4500 m C. effusa (Kunth) Steud. (syn.: Deyeuxia araeantha Pilg., Anthoxanthum L. D. effusa Kunth; C.fimckii Steud.); CO, VE, EC; 3000A. odoratum L.; CR, CO, VE, EC, PE; 2500-4000 m [Note: 4500m Native lo Europe, but introduced and naturalized in C. eminens (J.Presl) Steud. var. eminens (syn.: Deyeuxia páramo regions.] eminens J.Presl); CO, PE; 3600-3800 m C.fibrovaginata Lregaard [syn.: C. coarctata (Kunth) Steud. Aphanelytrum Hack. ( 1840) non Torre y ex Ea ton ( 1829); Deyeuxia coarctata A. procumbens Hack.; CO; 3000-4050 m Kunth, D.fuscata C.Presl]; CO, VE, EC, PE; 3650-4700 m C. guamanensis Escalona; EC; 4000-4400 m Aristida L. · C. heterophylla (Wedd.) Pilg. (syn.: Deyeuxia heterophyl/a A. enodis Hack.; PE; 2750-3100 m Wedd.; C. mül/eri Luces); VE, EC, PE; 3100-4650 m C. hirta (Sodiro ex Mille) Lregaard; EC; 2900-3900 m Arthrostylidium Rupr. C. intermedia (J.Presl) Steud. (syn.: C. humboldtiana Steud.; A. ecuadorense Judz. & L.G.Clark; CO; 2900-3100 m Deyeuxia secunda Pilg.); CR, PA, CO, EC, PE; 31004700m Au/onemia Goudot C. involuta Swallen; CO; 3700-3800 m A. bogotensis L.G.Clark, Londoño & Kobayashi; CO; 3100C. jamesonii Steud. [syn.: Deyeuxia janzesonii (Steud.) 3750m Munro ex Wedd.]; CO, EC, PE; 3600-4850 m A. pumi/a L.G.Clark & Londoño; CO; 2900-3250 m C. killipii Swallen; CO, VE; 3550-4150 m A. robusta L.G.Clark & Londoño; CO, VE; 2800-3200 m C. 1/anganatensis Lregaard; EC; 3500-4000 m A. trianae (Munro) McClure; CO; 3050-3600 m C. ligulata (Kunth) Hitchc. [syn.: C. podoplzora (Pilg.) Pilg.; Bothrioch/oa Kuntze Deyeuxia ligulata Kunth]; CO, VE, EC, PE; 3700-4850 B. saccharoides (sw:) Rydb. var. saccharoides (syn.: Androm [Note: These two species are sometimes considered pogon saccharoides Sw.); CO, PE?; 2900-3300 m distinct.]
1999]
PARAMOS
C. maaophylla (Pilg.) Pilg. (syn.: Deyeuxia macrophylla Pilg.); CO EC, PE?; 3800-4000 m G. mo/lis Pilg. [syn.: Deyeuxia setiflora Wedd.; C. setiflora (Wedd.) Pilg.]; EC; 3700-4600 m C. nuda (Pilg.) Pilg.; CR, PA, CO, EC; 2800-4250 m C. pisinna Swallen; CO, VE; 4000-4300 m C. pittieri Hack.; CR; 3100-3550 m C. planifolia (Kunth) Trin. ex Steud. (syn.: Deyeuxia p/anifolia Kunth); CO, VE, EC; 3000-4400 m 1 C. ramonae Escalona; VE; 3950 m C. recta (Kunth) Trin. ex Steud. (syn.: Deyeuxia recta Kunth); CO, EC, PE; 2900-4850 m C. rigescens (C.Presl) Scribn. [syn.: Deyeuxia rigescens (C.Presl) Türpe]; EC, PE; 3800-4500 m C. rígida (Kunth) Trin. ex Steud. (syn.: Deyeuxia rígida Kunth); CR, EC, PE; 3800-3900 m C. spicigera (J.Presl) Steud. (syn.: Deyeuxia spicigera J.Presl); PE; 4200 m C. steyermarkii Swallen; EC; 3450-4150 m C. tarmensis Pilg. [syn.: Deyeuxia tarmensis (Pi1g.) Sodiro]; EC?, PE; 2600-4100 m C. teretifoliaLa:gaard; EC; 4300-4850 m C. vicunarum (Wedd.) Pilg. (syn.: Deyeuxia vicunarum Wedd.); PE; 3300-4100 m . C. weberbaueri Tovar [syn.: Deyeuxia weberbaueri (Tovar) Rúgo1o]; CO, EC; 4000-4500 m
Chusquea Kunth (syn.: Swallenoch/oa McC1ure) Ref.: Clark, 1989; Soderstrom & Calderón, 1978. C. amistadensis L.G.Clark, Davidse & R.P.EIIis; CR, PA; 3050-3500 m C. angustifolia (Soderstr. & C.E.Calderón) L.G.C1ark; CO, VE; 2900-3500 m C. /atifolia L.G.Clark; CO; 2950-3380 m C. lehmannii Pilg.; CO, EC; 3100-3600 m; includes subsp. farinosa L.G.Ciark & Londoño and subsp. /ehmannii C. neurophyl/a L.G.Ciark; PE; 3200 m C. scandens Kunth; CO, EC?; 3300-3600 m C. spencei Ernst; CO, VE; 2900-3900 m C. subtessellata Hitchc.; CR, PA; 3000-3800 m C. tessellata Munro (syn.: C. weberbaueri Pilg.); CO, VE, EC, PE; 2750-4350 m C. vulcanalis (Soderstr. & C.E.Calderón) L.G.Ciark; CR; 3270-3350 m
Cinna L. C. poiformis (Kunth) Scribn. & Merr.; CR, CO, VE; 34003950m
Cortaderia Stapf C. bífida Pilg. (syn.: C. aristata Pilg.); CR, CO, VE, EC, PE; 2300-4100 m C. co/umbiana (Pilg.) Pilg.; CO, VE; 2800-3800 m C. hapalotricha (Pilg.) Conert (syn.: C. scabriflora Swallen, C. pungens Swallen); CR, PA, CO, VE, EC, PE; 29004900 m C. nítida (Kunth) Pilg. (syn.: C. sodiroana Hack.); CO, VE, EC; 2400-4200 m C. sericantha (Steud.) Hitchc.; CO, EC, PE; 3300-4500 m
121
Dactylis L. D. glomerata L.; CR, CO, VE, EC, PE; 3200-3800 m [Note: Native to Europe, but introduced, cultivated, and naturalized in páramo regions; in sorne places it is used as forage.]
Danthonia DC. D. secundiflora J.Presl; CO, VE, EC, PE; 2500-4200 m Deyeuxia Clarion ex P.Beauv. (see Calamagrostis) Dissanthelium Trin. D. calycinum (J.Presl) Hitchc. [syn.: D. mathewsii (Ba11) R.C.Foster & L.B.Sm.]; PE; 3700-4000 m
Elymus L. E. cordilleranus Davidse & R.W.Pohl [syn.: Agropyron attenuatum (Kunth) Roem. & Schult.]; CR, CO, EC, PE; 3200-4000 m
Festuca L. Re f.: Alexeev, 1986; Briceño & Morillo, 1994; Tovar, 1972.
F. amplissima Rupr. ex Fourn.; CR; 3000-3500 m F. andicola Kunth; CO, EC; 3300-4500 m F. arundinacea Schreb.; VE; 3300 m [Note: Introduced and escaped in páramo regions; it is sometimes cultivated as forage.] F. asplundii E.B.Aiexeev; CO, EC; 3600-4600 m F. azucarica E.B.A1exeev; CO; 3700 m F. breviaristata Pi1g.; CO, EC, PE; 3450-5000 m F. cajamarcae Pi1g.; PE; 3400-4100 m F. chimborazensis E.B.A1exeev; EC; 3900-4200 m F. cleefiana E.B.A1exeev; CO; 3600 m F. colombiana E.B.Aiexeev; CO; 3750-3800 m F. coromotensis B.Briceño; VE; 3300 m F. cundinamarcae E.B.Alexeev; CO; 3550 m F. densipaniculata E.B.Aiexeev; EC; 4600 m F. divergens Tovar; PE; 3600 m F. dolichophylla J.Presl; CR, CO, EC; 3400-4500 m F. elviae B.Briceño; VE; 3100-3550 m F.ferreyrae Tovar; PE; 3400-3500 m F.fragilis (Luces) B.Briceño (syn.: Helleriafragilis Luces); VE; 3600-4500 m F. glumosa Hack. ex E.B.A1exeev; EC; 3600-4700 m F. glyceriantha Pilg.; PE; 3600-4200 m F. herrerae Davidse; CR, PA; 2000-3300 m F. horridula Pilg.; PE; 3300-4000 m F. inarticulata Pilg.; PE; 3600 m F.jamesonii E.B.Alexeev; EC; 4350-4700 m F. loricata (Griseb.) Pilg.; PE; 3600-3900 m F. megalura Nutt.; EC; 3700 m F. parciflora Swallen; EC; 4100 m F. peruviana Infantes; EC, PE; 4000-4100 m F. procera Kunth; CO, EC; 3300-4000 m F. reclina/a Swa11en; CO; 3500-4000 m F. rigescens (J.Presl.) Kunth; EC; 3500-3800 m F. rigidifolia Tovar; PE; 3350-3900 m F. sublimis Pilg.; CO, EC; 2900-4300 m F. subulifolia Benth.; CO, EC; 3350-4300 m
122
F. F. F. F.
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
.' lálamancensis Davidse; CR; 3200~3400 m tarmensis Pilg.; PE; 3400~3800 m tolucensis Kunth; CR, CO, VE; 2900-4400 m vaginalis (Benth.) La;gaard (syn.: Poa vaginalis Benth.); CO, EC; 3500-4700 m
Helleria E.Fourn. (see Festucafragilis) Hierochloe R.Br. H. davidsei R.W.Poh1; CR, VE; 3100~3500 m H. redolens (Vah1) Roem. & Schu1t.; EC, PE; 3200~3800 m Holcus L. H. lanatus L.; CR, PA, CO, VE, EC; 2900~3600 m [Note: Native to Euro pe, but introduced and escaped in páramo regions.]
[VOL. 84
N. arista/a (Munro) Hitchc. (syn.: Planotia steubelii Pilg.); CO, EC, PE; 3250-4300 m N. asymmetrica L.G.C1ark; EC; 3Q00~3400 m N. elata (Kunth) Pilg. [syn.: N. nobilis (Munro) Pi1g.; Planotia nobilis Munro); CO, EC; .3400~3600 m Njimbri/igulata L.G.Ciark subsp.jimbriligulata; EC; 31003850 m [Note: This species is found in Polylepis forest.] N. laegaardii L.G.Clark; EC; 3200~3500 m N nana L.G.Ciark; EC; 3150-4100 m N. rígida L.G.Clark; EC; 3200-4300 m N. vi/losa L.G.Ciark; EC; 3750-4200 m Ortachne Nees ex Steud. O. erectifolia (Swallen) Clayton [syn.: Muhlenbergia erectifolia Swallen; Lorenzochloa erectifo!ia (Swallen) Reeder & C.Reeder; Parodie/la erectifolia (Swallen) Reeder & C.Reeder]; CR, CO, VE, EC, PE; 3300-4350 m
Lorenzochloa Reeder & C.Reeder (see Ortachne erectifolia) Parodiel/a Reeder & C.Reeder (see Ortachne erectifolia) Melica L. M. scabra Kunth; CO, EC, PE;
2600~3500
m
Muhlenbergia Schreb. M. angustata (J.Pres1) Kunth; CO, EC, PE; 2800-4000 m M. cajamarcensis La;gaard & Sánchez Vega; PE; 3000~ 3600 m M. calcicola Swallen; CR; 3400~3820 m . M. cleefii La;gaard; CO; 3500-4300 m Mjlabel/ata Mez; CR; 3300~3800 m M ligularis (Hack.) Hitchc.; CO, VE, EC, PE; 3350-4200 m [Note: This species is cultivated as forage.] M. nigra Hitchc.; CR; 2300~3500 m M. palmirensis Grignon & La;gaard; EC; 3200~3400 m M. peruviana (P.Beauv.) Steud.; EC, PE; 3500-4200 m M. rígida (Kunth) Trin.; EC, PE; 2300~3600 m M venezuelae Luces; VE; 3000-3600 m Nassel/a Desv. Ref.: Barkworth, 1990. N. brachyphyl/a (Hitchc.) Barkworth (syn.: S tipa brachyphylla Hitchc.); CO, EC, PE; 3300~3800 m N depaupera/a (Pilg.) Barkworth (syn.: Stipa depaupera/a Pilg.); CO, VE, EC; 3000-4200 m N. ibarrensis (Kunth) La;gaard (syn.: Stipa ibarrensis Kunth); EC; 2400~3200 m N. mexicana (Hitchc.) R. W.Poh1 (syn.: S tipa mexicana Hitchc.); CO, VE, PE; 3000-4100 m N mucronata (Kunth) R.W.Poh1 (syn.: Stipa mucronata Kunth); EC, VE, PE; 2900-4200 m N pubijlora (Trin. & Rupr.) Desv.; CO;EC, PE; 3000-3760 m Neurolepis Meisn. (syn.: Planotia Munro, nom. illeg.; ?latania Kunth, nom. illeg.) Ref.: Clark, 1996. N acuminatissima (Munro) Pi1g. (syn.: Planotia acuminatissima Munro); CO, EC; 2750~3300 m N aperta (Munro) Pi1g. (syn.: Planotia aperta Munro); CO, EC?; 3200~3600 m
Paspalum L. P. bonplandianum Flüggé; CO, VE, EC, PE; P. hirtum Kunth; CO; 3200~3800 m P. pilgerianum Chase; EC; 3400~3600 m P. prostratum Scribn. & Merr.; VE; 4000 m P. pygmaeum Hack.; VE; 3000~3600 m P. trianae Pilg.; CO; 2750-4000 m P. tuberosum Mez; EC, PE; 3000~3700 m
3000~3900
m
Phalaris L. P. aquatica L. (syn.: P. stenoptera Hack.); EC; 3400~3600 m [Note: Introduced and escaped in disturbed aquatic si tes in páramo regions; it is sometimes used as forage.] P. arundinacea L.; EC; 3240 m [Note: Introduced and escaped in páramo regions.] Piptochaetium J.Presl Ref.: Sánchez Vega, 1991. P. panicoides (Lam.) Desv.; CO, VE, EC, PE; 2900-4000 m P. sagasteguii Sánchez Vega; PE; 2750~3750 m P. tovarii Sánchez Vega; PE; 2800~3450 m; includes subsp. pilosa Sánchez Vega and subsp. tovarii P. tuberculatum Desv.; EC; 3400 m Poa L. P. annua L.; CR, CO, VE, EC, PE; 3500~4600 m [Note: Introduced from Europe.] P. boxiana Luces; VE; 2650~3600 m P. candamoana Pilg.; CO, EC; 3600-4300 m P. chamaeclinos Pilg.; EC, PE; 4000-4600 m P. chirripoensis R.W.Pohl; CR; 3500~3820 m P. cucullata Hack.; EC; 4200-4800 m P. horridula Pilg.; CO, EC, PE; 3600-4000 m P. leioclada Hack.; EC; 3100~3700 m P. mucuchachensis Luces; VE; 3400-3600 m P. orthophylla Pilg.; CO, VE; 3500-4600 m P. paramoensis La;gaard; EC; 3300-4500 m P. pardoana Pilg.; CO, EC, PE; 3550-4200 m
1999]
PARAMOS
P. paucijlora Roem. & Schult.; CO, VE, EC, PE; 3600-4800 m [Note: Sorne consider P. pardoana and P. paucijlora the same species.] P. petrosa Swallen; VE; 3350-4300 m P. pratensis L.; CR, CO, EC; 2500-3300 m [Note: Introduced from Europe, cultivated as forage, and sometimes escaped in páramo regions.] P. spicigera Tovar; PE; 3600 m P. slfbspicata (J.Presl) Kunth; CO, VE, EC, PE; 3000-4700 m P. talamancae R.W.Pohl; CR; 3300-3500 m P. trachyphylla Pilg.; CO, VE; 3300-4400 m P. trivialis L.; CO, EC; 4100-4500 m [Note: Introduced from Europe.] Poidium Nees P. monandrum (Hack.) Matthei [syn.: Briza monandra (Hack.) Pilg., B. mandoniana (Griseb.) Henr.]; EC, PE; 2900-3800 m Polypogon Desf. P. elongatus Kunth; CO, VE; 2500-4000 m P. interruptus Kunth; CO, EC, PE; 3500-4100 m Schizachyrium Nees S. hirtijlorum Nees; PE; 2700-3100 m Sporobolus R.Br. S. bogotensis Swallen & García-Barr.; CO; 2900-3100 m , S. lasiophyllus Pilg.; CO, VE, EC, PE; 2900-3700 m
123
V. myuros (L.) C.C.Gmel. [syn.: Festuca myuros L.; V. mega/ura (Nutt.) Rydb.]; CO, VE, EC, PE; 2800-4600 m; includes var. hirsuta Hack. and var. myuros
POLYGALACEAE Monnina Ruiz & Pav. M. aestuans (L.f.) DC.; CO; 3150-3550 m M. andreana Chodat; CO; 2800 m M. arborescens Ferreyra; CO; 3075-3600 m M. cf. bracteata Chodat; CO, VE; 3500-3650 m M. conferta Ruiz & Pav.; PE; 2800-4500 m M. confusa Ferreyra; PE; 3400-3550 m M crassifolia (Bonpl.) Kunth; CO, EC; 3500-4100 m M. densa Planch. & Lind.; CO, VE; 3200-3900 m [Note: This species is sometimes found growing on limestone outcrops.] M. e/ongata Planch. & Lind. ex Triana & Planch.; CO; 3300 m M involuta Ferreyra; CO; 3750 m M meridensis Planch. & Lind. ex Wedd.; VE; 2200-4000 m M. myrtilloides (Bonpl.) DC.; PE; 3500 m M. obtusifolia Kunth; CO, EC; 2750-3600 m M. pseudoaestuans Ferreyra & Wurdack; EC; 3350-3500 m M. revoluta Kunth; CO, EC; 3400-4200 m M sa/icifo/ia Ruiz & Pav. (syn.: M. nemorosa Kunth); CO, EC, PE; 3200-3750 m M. sandemanii Ferreyra; PE; 2900-3500 m
POLYGONACEAE
Swallenochloa McClure (see Chusquea)
Ref.: Brandbyge, 1989. Mueh/enbeckia Meisn. M. andina Brandbyge; EC; 2000-3950 m M. nummularia H.Gross.; PE; 3100-3400 m M. tamnifo/ia (Kunth) Meisn.; CR?, CO, VE, EC, PE; 33004000 m M. va/canica (Benth.) Endl.; CR?, CO, VE, EC, PE; 34004500m
Triniochloa Hitchc. T. andina Luces; CO, VE; 3520-3600 m T. stipoides (Kunth) Hitchc.; CR, CO, VE, EC; 2800-3600 m
Polygonum L. P. nepalense Meisn.; CO; 2800-4000 m [Note: This species is native to Asia and tropical Africa.]
Trisetum Pers. T.foliosum Swallen; CO, VE; 3500-3900 m T. irazuense (Kuntze) Hitchc.; CR, PA, CO, VE, EC; 19004200m T. oreophilum Louis-Marie; EC; 3600 m T. pringlei (Scribn. ex Beal) Hitchc.; CR; 2800-3800 m T. spicatum (L.) K.Richt. (syn.: T. andinum Benth.); CO, VE, EC, PE; 3900-4700m [Note: This species is introduced.] T. tonduzii Hitchc.; CR; 3100-3500 m
RumexL. R. acetosel/a L.; CR?, CO, VE, EC, PE; 3000-4550 m [Note: This species is native to Europe.] R. andinus Rech.f.; EC; 2700-3800 m R. /eptocau/is Brandbyge & Rech.f.; EC; 3500-3600 m R. obtusifo/ius L.; EC; 2250-3750 m [Note: This species is native to Europe.] R. peruanus Rech.f.; PE; 3400-3900 m R. tolimensis Wedd.; CO, EC; 3000-4500 m
Vulpia C.C.Gmel. V. australis (Nees ex Steud.) C.H.Blom (syn.: Festuca australis Nees ex Steud.); CO, VE, EC, PE; 3300-4100 m V bromoides (L.) Gray; CR, CO, EC, PE; 3250-4000 m V dertonensis (All.) Gola [syn.: Festuca dertonensis (All.) Asch. & Graebn.]; EC; 3300-4700 m
Calandrinia Kunth C. acau/is Kunth; CR, CO, VE, EC, PE; 3400-4700 m C. ciliata (Ruiz & Pav.) DC. (syn.: C. caulescens Kunth); CO, VE, EC; 2750-3600 m C. megarhiza Hemsl.; CR; 3500-3800 m
Stipa L. (see also synorryms under Nassel/a) S. hans-meyeri Pilg.; CR, EC, PE; 3400-4800 m S. ichu (Ruiz & Pav.) Kunth; CR, VE, EC, PE; 2500-4050 m S. inconspicua J.Presl; EC; 3500 m . S. mucronata Kunth; PE; 2700-3100 m
PORTULACACEAE
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MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
Mona O.Nilss. (see Montia meridensis) Montia L. (syn.: Mona O.Nilss.) Re f.: Lourteig, 1991. M. biapiculata Lourteig; CO; 3550--4150 m M. fontana L. (syn.: M. minar C.C.Gmei., M. pentandra Willd. ex Cham.); CO, EC, PE; 3150--4700 m M. meridensis Friedrich [syn.: Mona meridensis (Friedrich) O.Nilss.]; CO, VE; 3300--4850 m [Note: Steyermark & Rabe 97167 (US!) was misdetermined as Montia andina Rydb., a species ofMexico.]
POTAMOGETONACEAE Ref.: Haynes & Holm-Nielsen, in press.
Potamogeton L. P. illinoensis Morong; CO, EC, PE; 3375--4000 m P. paramoanus R.R.Haynes & Holm-Niels.; CO, VE, EC, PE; 3550--4100 m P. pusil/us L.; VE; 3550--4200 m Stuckenia Borner [syn.: Coleogeton (Rchb.) Les & R.R.Haynes] S. filiformis (Pers.) R.R.Haynes subsp. alpinus (Blytt) R.R.Haynes [syn.: Coleogetonjiliformis (Pers.) Les & R.R.Haynes subsp. alpinus (Blytt) Les & R.R.Haynes]; EC; 3200-3500 m
[VOL. 84
Ranunculus L. (syn.: Krapfia DC.; Rhopa/opodium O.E.Ulbr.) R. bonariensis Poir.; CO, VE; 3560--400qm R. crassirostratus Duncan var. cuchumatanensis Duncan; CR; 3500-3820 m R. cymba/aria Pursh; VE, EC; 4100--4200 m R.jlagel/iformis Sm.; CR, CO, VE, EC, PE; 1980--4300 m R. geranioides Kunth ex DC.; CO, VE, EC; ? m R. gigas Lourteig; PE; 4300--4800 m R. gusmanni Humb. ex Caldas [syn.: Krapfia gusmanni (Humb. ex Caldas) Stand!. & J.F.Macbr.; Rhopalopodium gusmanni (Humb. ex Caldas) O.E.Ulbr.]; CO, EC; 3900--4500 m R. krapfia DC. ex Deless. var. krapfia [syn.: Krapfia ranunculina DC.; Rhopalopodium ranunculinum (DC.) O.E.Ulbr.]; EC, PE; 3300--4300 m R. /ambayequensis T.Duncan & Ságast.; PE; 3300 m R. limosel/oides Turcz.; CO, VE, EC, PE; 3400--4300 m R. nubigenus Kunth ex DC.; CO, VE, EC; 2300--4200 m R. peruvianus Pers.; CR, PA, CO, EC, PE;3000--4400 m R. praemorsus Kunth ex DC. (syn.: R. argemonifolius Griseb.; CR, CO, VE, EC, PE; 2900--4500 m; includes var. amel!us (Briq.) T.Duncan, var. praemorsus, and var. sibbaldioides (Kunth ex DC.) Lourteig R. psychrophilus Wedd.; EC; 3450 m R. sandwithii Lourteig; CO; 3300-3350 m R. spaniophy!lus Lourteig; CO, VE; 3400--4400 m
PRIMULACEAE Lysimachia L. L. andina Sandwith; EC, PE; 2450-3700 m
PROTEACEAE
Rhopalopodium O.E.Ulbr. (see Ranunculus) Thalictrum L. T. podocarpum Kunth ex DC.; CO, VE; 3100-3400 m
Embothrium J.R.Forst. & G.Forst. (see Oreoca/lis)
RHAMNACEAE Oreocallis R.Br. (syn.: Embothrium J.R.Forst. & G.Forst.) O. grandiflora (Lam.) R.Br. (syn.: Embothrium grandiflorum Lam.); EC, PE; 2400-3900 m Roupala Aubl. R. pseudocordata Pittier; CO; 3050-3100 m
RANUNCULACEAE Ref.: Lourteig, 1956.
Anemone L. A.jamesonii Hook.; EC; 3250--4000 m Caltha L. C. sagittata Cav.; EC; 3400-5000 m Krapfia DC. (see Ranunculus) Laccopetalum Ulbr. L. giganteum (Wedd.) Ulbr. (Ranuncu/us giganteus Wedd.); PE; 4000--4500 m Oreithales Schltdl. O. integrifo/ia (DC.) Schltdl.; EC, PE; 3300--4300 m
Rhamnus L. [Note: A forest genus, but rarely found in subpáramo forest patches.] R. goudotiana Triana & Planch.; CO; 2800-3300 m R. oreodendron L.O.Williams; CR, PA; 2400-3500 m
ROSACEAE Ref.: Romoleroux, 1992, 1994, 1996.
Acaena Mutis ex L. .. A. m·gentea Ruiz & Pav.; EC, PE; 3100--4300 m A. cylindristachya Ruiz & Pav.; CR, CO, VE; 3200--4300 m A. e/ongata L.; CR, CO, VE, EC; 2000--4200 m A. ova/ifolia Ruiz & Pav.; CO, EC, PE; 2200-3900 m A. subincisa Wedd.; CO, EC; 3300--4550 m A. cf. torilicarpa Bitter; PE; 3200 m Alchemil/a L. (see Lachemilla) Aphanes L. A. cotopaxiensis Romoleroux & Frost-Olsen; EC; 38504100 m
1999]
PARAMOS
Fragaria L. F. vesca L.; CO, EC; 2800-3300 m [Note: Introduced from the Northern Hemisphere, cu1tivated and naturalized in páramo regions.] Geum L. G. peruvianum Focke; CO, EC, PE; 2500--4000 m Hesperomeles Lindl. (syn.: Osteomeles Lindl. as to Neotropics) H. cf. chiriquensis Woodson; PA; 2500-3400 m H. ferruginea (Pers.) Benth. (syn.: H. lanuginosa Ruiz & Pav. ex Hook., H. oblonga Lindl.); CO, VE, EC, PE; 2800-3900 m \H. goudotiana (Decnc.) Killip; CO; 2800--4000 m H. nítida Killip; CO; 3000 m H. obtusifolia (Pers.) Lindl. var. microphylla Wedd. (syn.: H. pernettyoides Wedd. var. microphylla Wedd.) in CO, EC at 3200--4200 m and var. obtusifolia [syn.: H. cuneata Lindl., H. glabrata Kunth, H.heterophylla (Ruiz & Pav. ex Lind.) Hook., H. obovata (Pittier) Standl.] in CR, PA, CO, VE, EC, PE at 2000--4000 m H. pachyphylla (Pittier) Killip (syn.: Osteomeles pachyphylla Pittier); CO; 2750-3600 m H. resinoso-punctata (Pittier) Pittier (syn.: Osteomeles resinoso-punctata Pittier); VE; 3800 m Holodiscus (K.Koch) Maxim. (see Sericotheca) Lachemilla (Focke) Rydb. Ref.: Gaviria, 1997; Perry, 1929; Rothmaler, 1935, 1937 L. andina (L.M.Perry) Rothm. (syn.: Alchemilla procumbens Rose var. andina L.M.Perry); CO, EC, PE; 3500--4500 m L. angustata Romoleroux; EC; 2850-3800 m L. aphanoides (Mutis ex L.f.) Rothm. (syn.: Alchemilla aphanoides Mutis ex L.f.); CR, PA, CO, VE, EC, PE; 2000--4500 m L. diplophylla (Diels) Rothm. (syn.: Alchemilla diplophylla Diels); EC; 4000--4180 m L. equisetiformis (Trev.) Rothm.; CO, VE; 3000--4600 m L. erodiifolia (Wedd.) Rothm. (syn.: Alchemilla erÓdiifolia Wedd.); EC; 4020 m L.fidvescens (L.M.Perry) Rothm. [syn.: Alchemilla venusta Schltdl. & Cham. var.fitlvescens L.M.Perry; L. venusta {Schltdl. & Cham.) Rydb.]; CO, VE, EC; 2250--4000 m L. galioides (Benth.) Rothm. (syn.: Alchemilla galioides Benth., A. radicans Rothm.); CO, EC, PE; 3000--4400 m L. hirta (L.M.Perry) Rothm. (syn.: Alchemilla procumbens Rose var. hirta L.M.Perry); CO, VE, EC; 2600--4400 m L. hispidula (L.M.Perry) Rothm. (syn.: Alchemilla hispidula L.M.Perry); CO, EC; 3000-5000 m L. holosericea (L.M.Perry) Rothm. (syn.: Alchemilla holosericea L.M.Perry); CO, EC; 3950--4400 m L. involucra/a Gaviria; VE; 3000-3200 m L.jamesonii (L.M.Perry) Rothm. (syn.: Alchemil/ajamesonii L.M.Perry); EC; 3350--4300 m L. killipii (Rothm.) Rothm. (syn.: Alchemilla killipii Rothm.); CO; 3100-3800 m L. loki-schmidtii Gaviria; VE; 3000-3200 m
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L. mandoniana (Wedd.) Rothm. [syn.: Alchemilla mandoniana Wedd., A. paludicola Rothm.; L. paludicola (Rothm.) Rothm.]; CO, VE, EC; 2900--4600 m L. moritziana Damm. [syn.: Alchemilla moritziana (Damm.) L.M.Perry]; CO, VE; 2800-3950 m L. nivalis (Kunth) Rothm. (syn.: Alchemilla nivalis Kunth); CO, VE, EC, PE; 2900--4700 m L. orbiculata (Ruiz & Pav.) Rydb. (syn.: Alchemilla orbiculata Ruiz & Pav.); CO, VE, EC, PE; 1600--4800 m L. pectina/a (Kunth) Rothm. (syn.: Alchemilla pectinata Kunth; L. pectinata subsp. caucana Rothm.); CR, PA, CO, EC; 2300--4000 m L. pelusa e Gaviria; VE; 4000--4600 m L. pinnata (Ruiz & Pav.) Rothm. (syn.: Alchemilla pinnata Ruiz & Pav.); CR, CO, VE, EC, PE; 3500--4300 m L. polylepis (Wedd.) Rothm. (syn.: Alchemilla polylepis Wedd.); CR, CO, VE; 3400--4300 m L. ramosissima (Rothm.) Rothm. (syn.: Alchemilla ramosissima Rothm.); VE; 3700--4300 m L. rivulorum (Rothm.) Rothm. (syn.: Alchemilla rivulorum Rothm., A. holmgrenii Rothm.); EC; 3800--4650 m L. rupestris (Kunth) Rothm. (syn.: Alchemilla rupestris Kunth); EC; 4000--4600 m L. sprucei (L.M.Perry) Rothm. (syn.: Alchemilla sprucei L.M.Perry); VE, EC; 3500--4100 m L. standleyi (L.M.Perry) Rothm. (syn.: Alchemilla standleyi L.M.Perry); CR; 2500-3800 m L. tanacetifolia Rothm.; CO, VE, EC, PE; 4000--4700 m L. trevirani Rothm. (syn.: L. lycopodioides Maguire); VE; 3150--4000 m L. tripartita (Ruiz & Pav.) Rydberg (syn.: Alchemilla tripartita Ruiz & Pav.); VE; 1500-3500 m L. unijlora Maguire; CO, EC; 3000--4400 m L. verticillata (Fielding & Gardner) Rothm. (syn.: Alchemilla verticillata Fielding & Gardner, A. resupinata Rothm.; L. arborescens Rothm.); CR, CO, VE, PE; 3000--4300 m L. vulcanica (Schltdl. & Cham.) Rydb. (syn.: Alchemilla vulcanica Schltdl. & Cham.); CO, EC, PE; 3200--4400 m Osteomeles Lindl. (see Hesperomeles) Polylepis Ruiz & Pav. Ref.: Simpson, 1979. P. incana Kunth; EC; 3000--4300 m P. lanuginosa Kunth; EC; 2700-3700 m P. microphylla (Wedd.) Bitter; EC; 3400-3650 m P. pauta Hieron.; EC; 3500--4200 m P. quadrijuga Bitter (syn.: P. boyacensis Cuatrec.); CO; 3300--4100 m P. quindiensis Cuatrec.; CO; 3500-3600 m P. racemosa Ruiz & Pav.; PE; 2900-3800 m P. retículata Hieron.; EC; 3200--4300 m P. sericea Wedd.; CO, VE, EC; 3200--4400 m P. weberbaueri Pilg.; EC, PE; 3000-3 700 m Poten tilla L. P. dombeyi Nestl. (syn.: P. andicola Benth.); EC; 3750--4500 m; includes var. andicola (Benth.) Lehm. and var. dombeyi P. heterosepala Fritsch; CR, CO, VE; 3000--4100 m
126
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
.' Rubits L. R. acanthophyllos Focke; CO, EC, PE; 3000-4000 m R. adenothallus Focke; EC, PE; 3000-3750 m , R. bogotensis Kunth (syn.: R. bogotensis subsp. eglandulosus Killip); CO, VE, EC; 2400-4100 m R. choachiensis Berger; CO; 3000-3600 m R. coriaceus Poir.; CO, EC; 3000-4300 m R. eriocarpus Liebm.; CR; 1000-3500 m R. gachetensis Berger; CO; 3100 m R. glabra tus Kurith; EC; 2400-4000 m R. /aegaardii Romoleroux; EC; 2700-3475 m R. loxensis Benth.; EC; 2600-3500 m R. macrocarpus Benth.; CO; 3300-3800 m R. megalococcus Focke; CO, EC, PE; 3000-3600 m R. nubigenus Kunth; CO, EC, PE; 2500-4200 m R. weberbaueri Focke; PE; 3400-3550 m Sericotheca Raf. [syn.: Ho/odiscus (K.Koch) Maxim.] S. argentea (L. f.) Rydb. [syn.: Spiraea argentea L.f.; Holodiscus argenteus (L.f) Maxim.]; CR, CO; 3000-4100
m
[VOL. 84
G. hypocarpium (L.) Endl. ex Griseb. subsp. hypocarpium [syn.: Relbunium hypocarpium (L.) Hemsl., R. croceum (Ruiz & Pav.) K.Schum., R. nitidum (Kunth) K.Schum.]; CR, PA, CO, VE, EC, PE; 3000-4000 m G. obovatum Kunth; EC; 2450-3750 m G. pseudotriflorum Dempster & Ehrend.; CO, EC, PE; 32003500 m G. pumilio Standl.; EC; 4200-4500 m G. uncinulatum DC.; CR; 3100-3800 m G. wedermanii Stand!.; EC; 3850 m. Manettia Mutis ex L. M. pichinchensis Wernham; EC; 3000-4200 m M. cf. trianae Wernham; CO; 2600-3350 m Nertera Banks & Sol. ex Gaertn. N granadensis (Mutis ex L. f.) Druce (syn.: N depressa Banks & Sol. ex Gaertn.); CR, CO, EC, PE; 3200-4350 m N tetrasperma Kunth; CO; 3200 m Relbunium (Endl.) Hook.f. (see Galium)
RUBIACEAE Arcytophyllum Willd. ex Schult. & Schult.f. Ref.: Mena V., 1990. A. aristatum Stand!.; CO, EC; 3500-4200 m A. cachirense (H.Karst.) K.Schum.; CO; 2800 m A. capitatum (Benth.) K.Schum.; CO, EC; 3300-3750 m A. ci/iolatum Stand!.; EC, PE; 3900 m A. ericoides (Willd. exRoem. & Schult.) Stand!.; PE; 30003400 m A. filiforme (Ruiz & Pav.) Stand!. [syn.: A. confertum (Ruiz & Pav.) var. bryoides (Willd. ex Roem. & Schult.) Stand!.]; CO, EC, PE; 3500-4300 m A. /avarum K.Schum. ex Stand!.; CR, PA; 3300-3800 m A. muticum (Wedd.) Stand!.; CR, CO, VE; 3000-4100 m A. nitidum (Kunth) Schltdl. [syn.: A. caracasanum (Kunth) Stand!.]; CO, VE; 3000-4200 m A. rivetii Danguy & Cherm.; EC, PE; 3400-3600 m A. setosum (Ruiz & Pav.) Schltdl.; CO, EC, PE; 3200-4300 m A. thymifo/ium (Ruiz & Pav.) Stand!.; EC, PE; 3200-3900 m A. venezue/anum Steyerm.; VE; 3000-3200 m A. vernicosum Stand!.; EC; 3400-3800 m Borreria G.Mey. B. anthospermoides DC.; CO; 2600-3350 m Ga/ium L. [syn.: Relbunium (Endl.) Hook.f.] Ref.: Dempster, 1981, 1982, 1990. G. aparine L.; CO; 3250-3300 m [Note: Introduced from Europe.] G. ascendens Willd. ex Spreng. (syn.: G. cuat¡·ecasasii Stand!.); CO; 3200-4000 m G. canescens Kunth (syn.: G. trianae Wernham); CR, CO, VE, EC; 3300-4100 m G. corymbosum Ruiz & Pav. [syn.: Relbunium ci/iatum (Ruiz & Pav.) Hemsl., R. hirsutum (Ruiz & Pav.) K.Schum.]; CO, EC, PE; 3400-4500 m
Sherardia L. S. arvensis L.; CO, EC, PE; 2800-3600 m [Note: Introduced from Europe.]
SAXIFRAGACEAE Saxífraga L. S. magellanica Poir.; EC, PE; 3500-4600 m; includes var. magellanica and var. peruviana (Sternb.) J.F.Macbr.
SCROPHULARIACEAE Ref.: Díaz Miranda, 1988; Holmgren & Molau, 1984. Alonsoa Ruiz & Pav. A. angustifo/ia Astholm, ined.; EC; ca. 3600 m [Note: This species is not meant to be published herein.] A. integrifolia Edwin; PE; ? m A. /actea (Diels) Astholm, ined.; CR, CO, EC; 2700-3900 m [Note: This species is not meant to be published herein.] A. linariifolia (Kunth) Steud.; PE; 2700-3900 m A. meridionalis (L.f.) Kuntze; CR, CO, VE, EC, PE; 6003950m A. procumbens Ruiz & Pav.; PE; 3400-3600 m A. pubescens Astholm, ined.; PE; 2800 m [Note: This species is not meant to be published herein.] A. trichanthera Astholm, ined.; PE; 3000 m [Note: This species is not meant to be published herein.] Aragoa Kunth Ref.: Fernández Alonso, 1993, 1995. A. abietina Kunth; CO; 3200-3800 m A. abscondita Fern.Alonso; CO; 3400-4200 m A. castroviejoi Fern.Alonso; VE; 3300-4000 m A. cleefii Fern.Alonso; CO; 3300-4000 m A. corrugatifolia Fern.Aionso; CO; 3300-3700 m
1999)
PARAMOS
A. A. A. A. A.
cundinamarcensis Fern.Alonso; CO; 3100-3400 m cupressina Kunth; CO, VE; 2900-4000 m dugandii Romero; CO; 3000-3550 m hammenii Fern.A1onso; CO; 3400 m kogiorum Romero; CO; 3000-4000 m; includes subsp. kogiorum and subsp. sevi/lae Fern.Alonso A. lucidula S.F.Biake; VE; 3100-3900 m; includes subsp. !anata Fern.Alonso and subsp. lucidula A. lycopodioides Benth.; CO, VE; 2800-4100 m A. occidentalis Pennell; CO; 3000-3500 m; in eludes subsp. occidentalis and subsp. refracta Fern.Alonso A. parvijlora Fern.Aionso; CO; 3700-4000 m A. pĂŠrez-arbelaeziana Romero; CO; 3500-3750 m A. romeroi Fern.Alonso; CO; 2400-3000 m A. tamana Fern.Aionso; CO, VE; 3100-3300 m Bartsia L. Ref.: Molau, 1990. B. alba Molau; EC; 3800-4250 m B. crisafullii N.H.Holmgren subsp. crisafiillii; EC, PE; 33003900m B. dijfi1sa Benth.; PE; 3770-3900 m B.jlava Molau; EC, PE; 3300-3600 m; includes subsp.jlava and subsp. minar Molau B. glandulifera Molau; CO, VE; 3200-4400 m B. inaequalis Benth.; EC; 2300-4100 m; includes subsp. inaequa/is B. /anijlora Benth.; CO, VE, EC; 2500-4400 m B. laticrenata Benth.; CO, EC; 3600-4500 m B. melampyroides (Kunth) Benth.; EC, PE; 3200-4600 m B. mutica (Kunth) Benth. (syn.: B. patriciae N.H.Holmgren); EC, PE; 3100-3800 m B. orthocarpijlora Benth.; CO, EC; 3250-4300 m; includes subsp. orthocarpijlora and subsp. vi/losa Mola u B. patens Benth.; PE; 3350-4900 m B. pedicularioides Benth.; CO, VE, EC; 2800-4500 m B. pumila Benth.; EC; 3800-4400 m B. ramosa Molau; CO, EC, PE; 3250-4100 m B. santolinifolia (Kunth) Benth.; CO, VE; 3000-4100 m B. sericea Molau; PE; 3000-3900 m B. stricta (Kunth) Benth.; CO, VE, EC; 3500-4700 m B. tenuis Molau; PE; 3150-3700 m B. tomentosa Molau; PE; 3400-4600 m B. trichophylla Wedd.; PE; 3500-3800 m B. weberbaueri Diels; EC, PE; 3600-3900 m Ca/ceo/aria L. Ref.: Molau, 1988. C. adenanthera Molau subsp. adenanthera; EC; 3600-3800 m C. adenocalyx Molau; CO; 3000-3100 m C. argentea Kunth; EC, PE; 2950-3900 m C. austra/is (Molau) Molau; EC; 3300-3400 m C. ballotifolia Kraenzl.; PE; 3150-3700 m C. barbato Molau; PE; 3400-4200 m C. bicrenata Ruiz & Pav.; PE; 3'500-4250 m C. caespitosa Molau; PE; 3350-3600 m C. cajabambae Kraenzl.; PE; 3300-4300 m C. calycina Benth.; EC, PE; 3300-3700 m
127
C. colombiana Pennell; CO, EC; 3500-3850 m C. commutata Molau; EC;3900-4000 m C. comosa Pennell; EC, PE; 3350-4500 m; includes subsp. comosa and subsp. elegans Molau C. concava Molau; PE; 3650-3900 m C. connatifolia Pennell; PE; 3400 m C. crenata Lam.; CO, EC; 3400-4000 m C. cumbemayensis Molau; PE; 3450-3600 m C. dejlexa Ruiz & Pav. subsp. cunea/a Molau; PE; 23004000m C. ericoides Vahl; EC, PE; 3400-4400 m; includes subsp. ericoides and subsp. peruviana Molau C.ferruginea Cav.; EC; 3500-4300 m C. frondosa Molau; EC; 3350 m C.fitsca Pennell; CO, EC, PE; 3450-3500 m C. gaultherioides M()lau; PE; 3000-3500 m C. gossypina Benth.; EC; 3300-4750 m C. helianthemoides Kunth; EC; 3400-3750 m C. hirtiflora Pennell; CO; 3000-3900 m C. hispida Benth. subsp. acaulis Molau; PE; 3300-3750 m C. hyssopifo/ia Kunth; EC; 4000-4300 m C. incarum Kraenzl. subsp .. sanchezii Molau; PE; 31503800 m C. injlexa Ruiz & Pav.; PE; 2800-3500 m C. irazuensis Donn.Sm.; CR, PA; 2500-3550 m C. laevis Molau; PE; 3200-3550 m C. /amiifo/ia Kunth; CO, EC; 3500-4200 m C. leucanthera Edwin; CO, VE; 3250-4300 m C. linearis Ruiz & Pav.; PE; 3500-4350 m C. /ojensis Pennell: EC; 2900 m C. macula/a Edwin; PE; 1700-3600 m C. melissifolia Benth. subsp. melissifo/ia; PE; 3500-4000 m C. mexicana Benth. (syn.: C. palustris Sodiro ex Kraenzl.); CR?, CO, VE, EC, PE; 1200-4100 m; includes subsp. mexicana and subsp. perijensis (Pennell) Molau C. micans Molau; PE; 3200-3900 m C. microbefaria Kraenzl.; CR, CO, VE, EC; 2500-4300 m; includes subsp.fruticosa (Pennell) Molau, subsp. microbefarĂa, and subsp. tatamana (Pennell) Molau C. moyobambae Kraenzl.; PE; 3400-3600 m C. nevadensis (Pennell) Stand!. subsp. meridensis (Pennell) Molau; VE; 3000-4000 m C. nivalis Kunth subsp. cerasifo/ia (Benth.) Molau; EC, PE; 3300-3900 m C. obtusa Molau; EC; 3300 m C. penlandii Pennell; CO, EC; 3700-3750 m; includes subsp. penlandii and subsp. puraceensis (Pennell) Molau C. percaespitosa Wooden; PE; 3500-4300 m C. perfoliata L. f.; CR, PA, CO, VE, EC; 2400-3800 m C. phaceliifolia Edwin; PE; 3350-3 750 m C. phaeotricha Molau; EC; 3300-3700 m C. pilosa Molau; PE; 3350 m C. pumila Edwin; PE; 3150-3550 m C. reichlinii Edwin; PE; 3000-3900 m C. rhododendroides Kraenzl.; PE; 3400-3700 m C. rosmarinifolia Lam.; EC; 3400-3800 m C. salicifolia Ruiz & Pav. subsp. nigricans Molau; PE; 35003750 m
128
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
C. sibthorpioides Kunth; PE; 3850 m C. tetragona Benth.; PE; 3450-3900 m; inc1udes subsp. endopogon (Kraenzl.) Mo1au and subsp. tetragona C. !rilaba Edwin; PE; 3350--4300 m C. tri/abata Hemsl. (syn.: C. aequi/ateralis Edwin); CR, CO, VE, EC, PE; 2100-4100 m; includes subsp. aequilateralis (Edwin) Mo1au and subsp. tri/abata C. tripartita Ruiz & Pav.; CR, CO, VE, EC, PE; 1600--3650 m C. utricularioides Benth.; PE; 3400-4000 m C. virgata Ruiz & Pav.; EC, PE; 3500-4300 m C. viscosa Ruiz & Pav.; PE; 3700 m C. weberbaueriana Kraenzl.; PE; 3600-4300 m Castil/eja Mutis ex L.f. (syn.: Orthocarpus Nutt.) C. alpicola T.I.Chuang & Heckard; PE; 4000 m C. arvensis Sch1tdl. & Cham.; CO, VE?, EC; 2000-3800 m C. breviflora Benth.; CO, VE; 3800-4300 m C. cerroana Edwin; EC, PE; 3550-3900 m C. ecuadorensis N.H.Ho1mgren; EC; 3400-4400 m C.fissifolia L.f. [syn.: C.fissifolia subsp. divaricata (Benth.) Wedd., subsp.fi'uticosa Wedd. ex Melch., subsp. integrifo/ia (L. f.) Wedd.]; CR, CO, VE, EC, PE; 3000-4600 m C. irazuensis Oerst.; CR, PA; 1800-3820 m C. meridensis Pennell; VE; 4100-4400 m C. nubigena Kunth; EC, PE; 3600-4750 m C. profunda T.I.Chuang & Heckard; PE; 3300-3350 m C. pumila (Benth.) Wedd.; CO, EC, PE; 3600-4500 m C. quirosii Standl.; CR, PA; 3500-3800 m C. talamancensis N.H.Ho1mgren; CR, PA; 3100-3820 rn C. vadosa T.l.Chuang & Heckard (syn.: C. peruviana T.I.Chuang & Heckard); PE; 3400-4000 rn C. virgata (Wedd.) Edwin; CO, EC, PE; 2500-4650 m Digitalis L. D. purpurea L.; CO, EC; 2800-4000 m [Note: Introduced frorn Europe, cu1tivated in páramo regions asan herba1 and escaped.] Gratio/a L. G. bogotensis Cortés; CO?, EC; 2000-3500 m Lamourouxia Kunth L. sylvatica Benth.; EC, PE; 2800-4500 rn L. virgata Kunth; CO, EC; 3400-3600 rn Limase/la L. L. americana G!ück; CR?, VE; 3000-4000 m L. aquatica L.; CR, VE, EC; 4100 m L. australis R.Br.; CR, CO; 3500-4250 rn L. subulata E.Ives; VE, EC; 3700-4450 m Mimulus L. M. glabratus Kunth; CO, VE, EC, PE; 1800-4200 m Orthocarpus Nutt. (see Casti/leja) Ourisia Comm. ex Juss. O. chamaedrifo/ia Benth.; CO, VE, EC, PE; 3500-4300 m.
[VOL. 84
O. muscosa Benth. (syn.: O. nana Benth.); CO, EC, 3900-4600 m
PE~·
Pedicularis L. P. incurva Benth.; CO, EC; 3400-4300 m Sibthorpia L. S. repens (L.) Kuntze (syn.: S. pichinchensis Kunth); CR, CO, VE, EC, PE; 2200-4300 rn Veronica L.
V. arvensis L.; CR, CO, EC, PE; 3200-4200 m [Note: This species is native to Europe.]' V. peregrina L.; CO, EC, PE; 3350-4200 m; includes var. peregrina and var. xalapensis (Kunth)'St.John & Warren V. persica Poir.; CO, VE, EC, PE; 1700-3650 m [Note: This species is native to SW Asia.] V. serpyl/ifolia L.; CR, CO, VE; EC, PE; 2900-4300 m; ineludes var. humifitsa (Dickson) Vah! and var. selpyl/ifo/ia
SOLANACEAE Acnistus Schott (see Saracha) Cestrum L. C. buxifolium Kunth (syn.: C. melanochloranthum Duna!); CO, VE; 3000-4100 m C. microphyl/um Linden ex Duna!; CO; 2200-3400 m C. petiolare Kunth; VE; 2000-3830 m C. rigidifolium Francey; CO; 3500-4000 m Deprea Raf. D. paneroi Benitez & M.Martínez; VE; 2650-2900 m Hebecladus Miers (see Jaltomata viridiflora) Jaltomata Schltdl. J. paneroi Mione & S.Leiva; PE; 3200-3550 m J. viridiflora (Kunth) M.Nee & Mione [syn.: Hebecladus viridiflorus (Kunth) Miers]; CO, VE, EC; 2400-3400 m Nicotiana L. N. thyrsiflora Goodspeed; PE; 3300-4000 m Nierembergia Ruiz & Pav. N. repens Ruiz & Pav.; PE; 3650 rn N. spathulata Kunth; CO, EC; 2700-3050 m Salpichroa Miers S. diffusa Miers; CO, EC; 3100-4200 m S. glandulosa (Hook.) Miers; PE; 3050-4700 m S. tristis Miers var. tristis; CO, VE; 2800-3600 m Saracha Ruiz & Pav. S. punctata Ruiz & Pav. [syn.: Acnistus /indenianus (Miers) Hunz.; S. ovala (Miers) Hunz.]; CO, VE, EC; 3050--3800 m S. quitensis (Hook.) Miers [syn.: Acnistus quitensis (Hook.) Hunz.]; CO, VE, EC, PE; 3000-4000 m
1999]
PARAMOS
Solanum L. S. albicans (Ochoa) Ochoa (syn.: S. acaule Bitter var. a/bicans Ochoa); PE; 3500 m S andreanum Baker (syn.: S. sujfrutescens Correll); CO, EC; 2800-3800 m S. barbulatum Zah1br. (syn.: S. ecuadorense Bitter); CO, EC, PE; 2500-4000 m S. callianthum C.V.Morton; CO; 2000-3500 m S. chíquidenum Ochoa; PE; 2800-3700 m; inc1udes var. chiquidenum and var. porconense Ochoa S. chomatophilum Bitter; EC, PE; 3350-4000 m; includes var. angustifolium Correll and var. chomatophilum S. clivorum S.Knapp; PE; 3000-3200 m [Note: Weedy species found along páramo margins.] S. co/ombianum Duna! (syn.: S. do/ichocarpum Bitter, S. filamentum Correll); CO, VE?; 3000-3600 m S. cornifo/ium Duna!; CO, VE, EC; 2300-3000 m [Note: Weedy species found along páramo margins.] S. cutervanum Zahlbr. (syn.: S. pu/veru/entum Pers.); PE; 2600-3600 m S. deflexiflorum Bitter; CO;? m [Note: Weedy species found along páramo margins.] S. donachui (Ochoa) Ochoa; CO; 3200-3260 m S.jlahaultii Bitter; CO, VE; 3300-3700 m S. foetens S.Knapp; VE; ? m [Note: Weedy species found along páramo margins.] S.fragi/e Wedd.; PE; 3400-3600 m S. garcia-barrigae Ochoa; CO; 3900 m S. goniocaulon S.Knapp; EC, PE; 2000-3000 m [Note: Weedy species found along páramo margins.] S. hypa/eurotrichum Bitter; EC; 2000-3500 m S.ja/cae Ochoa; PE; 3100-3500 m; includes var.jalcae and var. pubescens Correll S. /aurifrons Bitter; CO; 2800-3000 m [Note: Weedy species found along páramo margins.] S. /eiophyllum Benth. (syn.: S. benthamii Duna!); EC; 34003650 m S. matureca/vans Bitter; EC, PE; 2000-3000 m [Note: Weedy species found along páramo margins.] S. moscopanum Hawkes; CO; 2900-3400 m [Note: This species is found in cultivated fields and páramo pasturelands.] S. nigrescens M.Martens & Galeotti; CR?, CO; 1800---3400 m S. nitidum Ruiz & Pav.; EC, PE; 3500-4300 m S. nutans Ruiz & Pav.; CO, EC, PE; 2000-3500 m S. oblongifo/ium Duna] (syn.: S. jamesonii Bitter, S. hypomalacophylum Bitter); CO, VE, EC; 2950-3700 m S. ochranthum Duna]; CO; 3300 m S. paramoense Bitter; VE; 3300 m S. paucijugum Bitter; EC; 3200-4100 m S. psychotrioides Duna]; CO; 3150-3450 m S. solisii Hawkes; EC; 3500-4000 m S. stellatig/andulosum Bitter; CO; 2700-3700 m S. stenophyllum Duna] (syn.: S. bogotense Duna!, S. neriifolium Bitter, S. toliménse Wedd.); CO, EC; 27004000m S. storkii C.V.Morton & Standl.; CR; 2800-3500 m S. subpanduratum Ochoa; VE; 3400-3600 m
129
S. tuberosum L. (syn.: S. andigena Juz. & Bukasov); CO; 3300-3 700 m; includes subsp. andigena (Juz. & Bukasov) Hawkes and subsp. tuberosum [Note: Introduced as a crop p Jant.] S. tuquerrense Hawkes; EC; 3000-3600 m S. vacciniiflorum Standl. & L.O.Williams; CR; ? m [Note: Weedy species found along páramo margins.] S. validinervium Benitez & S.Knapp; VE; 2000-3000 m [Note: Weedy species found along páramo margins.] S. venosum Duna!; CO, EC; 2000-3500 m S. youngii S.Knapp; PE; 2800-3500 m S. zah/bruckneri Bitter; EC; 3400-3 800 m
SYMPLOCACEAE Symplocos Jacq. Ref.: Stahl, 1991, 1993. S. fimbria/a B.Stahl; EC, PE; 3400-3700 m S. incahuasensis Sagást. & M.O.Dillon; PE; 3300-3900 m S. nana Brand; EC, PE; 3200-4000 m S. niva/is Linden ex Brand; CO; 3600-3700 m S. oreophila Almeda; CR; 3150 m S. quitensis Brand; EC; 2900-4000 m S. rejlexa A.DC.; EC, PE; 2700-3550 m S. scabra J.F.Macbr.; PE; 3350-3600 m S. tamana Steyerrn.; VE; 3300-3500 m S. theiformis (L. f.) Oken; CO; 3000-3450 m
THEACEAE Ternstroemia Mutis ex L.f. T. camel/iaefolia Lind. & Planch.; CO; 2800 m T. meridionalis Mutis ex L. f.; CO; 2600-3575 m
TROPAEOLACEAE Tropaeolum L. T. tuberosum Ruiz & Pav. subsp. silvestre Sparre; CO; 35003650 m [Note: The subsp. tuberosum is cultivated as a tuber plant.]
URTICACEAE Parietaria L. P. debilis G.Forst.; CO; 3750 m Pi/ea Lindl. [Note: A forest genus, but rarely found in isolated páramo forest patches.] P.fal/ax Wedd.; CO; 1900-3700 m P. pulegifolia (Poir.) Wedd.; PE; 3100-3600 m P. serpyllacea (Kunth) Liebm.; EC; 1800-3700 m Urtica L. U ballotifolia Wedd.; CO, VE; 3600-3900 m; perhaps equal to U leptophylla Kunth U echinata Benth.; EC; 4050-4150 m U jlabellata Kunth; EC; 4000-4050 m U /eptophylla Kunth; CO, VE; 3300-4100 m
130_
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
VALERIANACEAE Re f.: Eriksen, 1989a, 1989b; Xen a de Enrech; 1992. Belonanthus Graebn. B. longitubulosus Schmale; PE; 4000-4100 m [Note: Belonanthus was placed within Valeriana as a section within sub gen. Phyl!actis (Eriksen, 1989a); however, the combinations at the species leve! have not yet been made in Valeriana and Ido not intend todo it here.] Valeriana L. [syn.: Aretiastrum (DC.) Spach., Belonanthus Graebn., Phy!lactis Pers., Stangea Graebn.; but see also Belonanthus longitubulosus] V. adscendens Turcz.; CO, EC, PE; 3250-4500 m V. alypifolia Kunth (syn.: V. hartwegiana Briq.); EC; 3800:-' 4900 m; includes subsp. alypifo/ia and subsp. argen(e¿marginata B.Eriksen V. arborea Killip & Cuatrec.; CO; 3650-4350 m V. aretioides Kunth; EC; 3700-4600 m V. bracteata Benth. ( syn.: V. ligulifolia Killip ); CO; EC, PE; 3250-4400 m V. bractescens (Hook.) Hóck (syn.: V. meridana Briq.); VE; 3100-4100 m V. buxifolia F.G.Mey.; EC; 3500-3800 m V. cernua B.Eriksen; EC; 3800-3900 m V. cerosifolia Xena; VE; 2850--3300 m V. clematitis Kunth (syn.: V. flexuosa Graebn.); CO, EC; 3300-4000 m V. comosa B.Eriksen; PE; 3900 m V. connata Ruiz & Pav.; PE; 4000-4200 m V. convallarioides (Schmale) B.B.Larsen; EC, PE; 34003500m V. crassifolia Kunth; CO; 3400-4150 m V. cuatrecasasii F.G.Mey.; CO; 2800 m V. cumbemayensis B.Eriksen; PE; 3500-3600 m V. decussata Ruiz & Pav. subsp. po/emonioides (Kunth) B.Eriksen (syn.: V. polemonoides Kunth, V. clematoides Graebn.); CO, EC, PE; 2650-3600 m V. engleriana Hóck; CO; 3850 m V. granataea Xena; VE; 3300-3600 m V. henrici (Graebn.) B.Erjksen; EC; 4200-4400 m V. hirte/la Kunth; EC; 2900-3700 m V. humboldtii Hook. & Arn.; PE; 3400-3560 m V. imbricata Killip; EC, PE; 3200-3450 m V. interrupto Ruiz & Pav.; PE; 3150-3600 m V. karstenii Briq.; CO; 3400-4300 m V. microphylla Kunth (syn.: V. amphi/ophis Graebn., V. alophis Graebn., V. bonplandiana Wedd., V. hieronymii Graebn., V. revoluta Diels.); CO, EC, PE; 2700-4700 m V. niphobia Briq.; EC, PE; 3300-4650 m V. parviflora (Trevir.) Hóck; VE; 3000-4400 m V. phylicoides (Turcz.) Briq.; VE; 3100-4300 m V. pilosa Ruiz & Pav. (syn.: V. longifolia Kunth); CR, CO, EC, PE; 3100-4500 m V. plantaginea Kunth; CO, VE, EC, PE; 3200-4600 m V. prionophylla Stand!.; CR, PA; 3300-3800 m V. pulchella M.Martens & Galeotti; CR, PA; 3350-3800 m V. pyramidalis Kunth; EC; 2000-4000 m
[VOL. 84
V. quindiensis Killip; CO; 3400-4000 m V. qu{rorana Xena (!>yn.: V. foliosa Pittier & ~illip non Phi!.); VE; 3200 m V. rigida Ruiz & Pav. [syn.: Phyllactis rigida (Ruiz & Pav.) Pers.]; CO, EC, PE; 3300-4700 m V. robertanifolia Briq.; VE; 4000 m V. rosaliana F.G.Mey.; VE; 3000-3400 m V. rumicoides Wedd.; CO, EC; 3350-4100 m V. secunda B.Eriksen; EC; 3800-4400 m V. smithii Killip; CO; 3750-3850 m V. spatulata Ruiz & Pav. [syn.: Belonanthus spatulatus (Ruiz & Pav.) Schmale]; PE; 4650 m V. spicata (Turcz.) Briq.; VE; 3300-4400 m V. spirojlora B.B.Larsen; CO; 3600-3800 m V. stenophylla Killip; CO; 3300-3800 m V. tachirensis Xena; VE; 3000-3300 m V. tatamana Killip; CO, EC; 2900-4100 ni. V. thalictroides Graebn.; PE; 3600 m V. triphyl/a Kunth [syn.: V. mutisiana (Wedd.) Hóck]; CO, VE; 2800-4100 m V. triplinervis (Turcz.) Briq.; VE; 3000-3500 m V. venezuelana Briq.; VE; 2500-3600 m V. vetasana Killip; CO; 3450-3900 m V. weberbaueri Graebn.; PE; 3650 m
VERBENACEAE Hierobotana Briq. H. injlata (Kunth) Briq.; EC; 2450-3700 m Lippia L. L. schlimii Turcz. var. glabrescens Moldenke; CO; 1650-
3300m Verbena L. V. glabrata Kunth; EC; 3450 m V. microphylla Kunth; EC; 3700 m V. occulta Moldenke; PE; 4000 m
VIOLACEAE Hybanthus Jacq. H. parvijlorus (Mutis ex L.f.) Baill. (syn.: Viola venezuelensis Steyerm.); CO, VE, EC, PE; 2800-3600 m Viola L. V. adenophylla Killip; EC; 3400 m V. arguta Kunth var. meridionalis (W.Becker) L.B.Sm. &
A.Fernández; CO, EC, PE; 3100-3450 m V. bangii Rusby (syn.: V. nivalis Benth. non Roem. &
Schult.); EC, PE; 3900-4600 m V. glandulifera Hook.; EC, PE; 3300-4350 m V. humboldtii Triana & Planch.; CO, EC; 2400-3350 m V. humi!is Kunth; CO; 3200-3800 m V. membranacea W.Becker; PE; 3300-4000 m V. nannei Polakowsky; CR; 1500-3500 m V. polycephala H.E.Ballard & P.J0rg.; EC; 3800-4700 m [Note: New name for V. parvifolia Benth. (nom. illeg.), see Novon 7: 13. 1997.]
1999]
.1
~· \
! r 1
PARAMOS
131
Badillo, V. M. 1994 [1995]. Enumeración de las Compuestas (Asteraceae) de Venezuela. Revista Fac. Agron. (Univ. Central Venez.) 45: 1-191. - - - . 1997. Los géneros d~ la~ Compositae (Asteraceae) de Venezuela: Clave artificml para su determinación · Ernstia 6: 51-168. Balslev, H. 1996. Juncaceae. Fl. Neotrop. Monogr. 68: 1-168. ---.&R. Briones: 1982. Las Gentianáceas del Parque Nacwnal Cotopaxi, Ecuador. Pub!. Mus. Ci. Nat. Ecuador, Ser. Revista 3(3): 12-19. Barkworth, M. E. 1990. Nassella (Gramineae, Stipeae): Revised interpretation and nomenclatura! changes. Taxon 39: 597-614. Berry, P. E. 1985. The systematics ofthe apefalous fuchsias VlSCACEAE of South America, Fuchsia sect. Hemsleyella (OnaRef.: Kuijt, 1986. graceae). Ann. Missouri Bot. Gard. 72: 213-251. Bishop, L. E. 1978. Revision of the genus Cochlidium Dendrophthora Eichler (Grammitidaceae). Amer. Fem J. 68: 76-94. D. ambigua Kuijt; CR, CO, EC; 3250-3400 m - - - . 1988. Ceradenia, a new genus ofGrammltidaceae. D. chrysostachya (J.Presl) Urb.; CO; 2650-3600 m Amer. Fem J. 78: 1-5. D. clavata (Benth.) Urb.; CO, VE, EC; 3100-3700 m - - - . 1989. Zygophlebia, a new genus of GrammiD. costaricensis Urb.; CR; 3100-3500 m tidaceae. Amer. Fem J. 79: 103-118. D. densifolia Kuijt; PE; 3450 m - - - & A. R. Smith. 1992. Revision of the fem genus D. lindeniana Tiegh.; CO, VE, EC; 2850-3300 m Enterosora (Grammitidaceae) in the New World. Syst. D. longepedunculata Rizzini; VE; 2600-3400 m Bot. 17: 345-362. D. paucifolia (Rusby) Kuijt; CO; 3100-3200 m Bohlin, J.-E. 1988. A monograph ofthe genus Colignonia D. pearcei (Rus by) Kuijt; CO; 2700-3000 m (Nyctaginaceae). Nordic J. Bot. 8:231-252. D. squamigera (Benth.) Kuntze (syn.: D. biserrula Eichler); Brako, L. & J. Zarucchi (eds.). 1993. Catalogue ofthe flowering plants and gymnosperms of Peru [Catálogo de las CR, P A, CO, VE, EC; 2800~000 m angiospermas y gimnospermas del Perú]. Mono gr. Syst. Bot. Missouri Bot. Gard. 45: i-xl, 1-1286. WINTERACEAE Brandbyge, J. 1989. Polygonaceae. Fl. Ecuador 38: 1-62. Bremer, K. (ed.). 1994. Asteraceae: Cladistics and classifiDrimys J.R.Forst. & G.Forst. cation. Timber Press, Portland, Oregon. [Note: A forest genus, but rarely found in subpáramo Briceño, B. & G. Morillo. 1994. El género Festuca L. forest patches.] (Poaceae) en los páramos de Mérida. Emstia 4: 73-88. D. granadensis L.f. var. grandijlora Hieran.; CR, PA, CO; Bywater, M. & G. E. Wickens. 1984. New World species of 3100-3700 m the genus Crassula. Kew Bull. 39: 699-728. Cabrera, A. L. 1965. Revisión del género Mutisia (Compositae). Opera Lilloana 13: 1-227. XYRIDACEAE Chung, I. 1965. Revision of Barnadesia (CompositaeMutisieae). Privately published, Chicago. Xyris L. Clark, L. G. 1989. Systematics ofChusquea, sect. SwallenoX subulata Ruiz & Pav. [syn.: X acutifolia (Heimerl) chloa, sect. Verticillata, sect. Serpentes and sect. LongiMalme]; CR, PA, CO, VE, EC, PE; 3000~250 m; infolia (Poaceae, Bambusoideae). Syst. Bot. Monogr. 27: eludes var. acutifolia Heimerl, var. breviscapa Idrobo & 1-127. L.B.Sm., and var. subulata - - - . 1996. Six new species of Neurolepis (Poaceae: Bambusoideae: Bambuseae) from Ecuador and Peru. Novan 6: 335-350. Cleef, A. M. 1981. The vegetation of the páramos of the Colombian Cordillera Oriental. Diss. Bot. 61: 1-320. J. Cramer, Vaduz Literature Cited in Checklist ofVascular Plants Clemants, S. E. 1995. Be)aria. Pp. 54-106. In: Luteyn (ed.), Alexeev, E. B. 1986. Festuca L. (Poaceae) in Venezuela, 1995. Colombia et Ecuador. Novosti Sistematiki Vyschnij Cook, C. D. K. ·1985. A revision of the genus Efod ea Rastenii 23: 5-23. (Hydrocharitaceae). Aquatic Bot. 21: 111-156. Allen, C. K. 1933. A monograph ofthe American species of Crane, E. H. 1997 [1998]. A revised circumscription ofthe the genus Halenia. Ann. Missouri Bot. Gard. 20: 119-222. genera of the fern family Vittariaceae. Syst. Bot. 22: Anderberg, A. A. 1991. Taxonomy and phylogeny of the 509-518. tribe Gnaphalieae (Asteraceae). Opera Bot. 104: 1-195. Cronquist, A. 1981. An integrated system of classification of Aristeguieta, L. 1964. Compositae. Fl. Venezuela 10(1, 2): flowering plants. Columbia University Press, New York. 1-941. Cruden, R. 1991. A revision of Jsidrogalvia (Liliaceae): Ayers, T. J. 1997. Three new speceis of Lysipomia (LobeliRecognition for Ruiz and Pavón's genus. Syst. Bot. 16: aceae) endemic to the páramos of southern Ecuador. 270-282. Brittonia 49: 433~40.
V. pusillima Wedd.; PE; 3550-3800 m V. pygmaea Juss. ex Poir.; EO, PE; 3600~400 m V. stipularis Sw. (syn.: V. obliquifolia Turcz.); VE; 24002837 m V. tricolor L.; EC; 2500-3500 m [Note: Introduced from Europe, cultivated, and sometimes escapes in páramo regions.] V. veronicifolia Planch. & Lindl. ex Triana & Planch.; CO; 2950-3700 m V. weberbaueri W.Becker; PE; 3150-3200 m
132
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
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OF
PARAMO
The following list of páramo localities (gazetteer) is organized first by country. Within each country (listed from north to south, except Costa Rica and Panama, which are given together) the locality names are given in strict alphabetical order omitting al! Spanish articles such as El, La, Las, and Los. The articles are given after the proper name, and when placed within parentheses their use is optional or variable. Localities with the same name are then designated as (1), (2), etc. The most commonly used name is detailed, while less-used names are cross-referenced to the primary name. The highest known altitude is given next, followed by latitude and longitude (rounded off to the nearest minute). The next two headings are called "Division 1" (i.e., primary political divisions such as department in Colombia and Peru, state in Venezuela, and province in Costa Rica, Panama, and Ecuador) and "Division 2" (i.e., secondary political divisions such as municipality in Colombia, district in Venezuela, canton in Costa Rica and Ecuador, and province in Peru), when these are known (Division 2 was not used at all for Ecuador). Ifthe páramo locality covers two or more political divisions within Division 1, these divisions are separated by a slash (/). If the páramo locality covers two or more political divisions within Division 2, the Division 2 divisions are separated by an ampersand (&) and the Division 1 by a slash (/). [Example 1: In Colombia, Cerro Agua Blanca overlaps the Municipality Puracé of the Departament Cauca and the Municipality La Argentina of the Department Huila; therefore, Di vis ion 1 reads "Cauca/Huila" and Division 2 reads "Puracé/La Argentina." Example 2: In Colombia, Cerro Animas overlaps the Municipalities ofLa Cruz andEl Tablón of the Department Nariño and the Municipality of Santa Rosa of the Department Cauca; therefore, Division 1 reads "Nariño/Cauca" and Division 2 reads "La Cruz & El Tablón/Santa Rosa."] The last entry, "Notes," usually gives more details about the geo-
LOCALITIES
graphical location, direction from a large or wellknown city, connection with other nearby páramos, other spellings for the primary name, cross-references to other names, and comments. The primary sources oflocality information for the páramo gazetteer (i.e., the names, elevations, latitude and longitude, reference points in "Notes," etc.) were individual country maps ( topographical, geopolitical, road maps, tourist maps, etc.), individual country gazetteers, label data from herbarium specimens, and literature references. The complete database includes detailed references for each individual entry, which are not included in this book but are available upon request. The main sources of data for each country are listed at the end of this chapter; the generalliterature references cited in the text are found in the bibliography at the end of the book. It was not always easy to determine the exact name, geographical position, or political division of a particular páramo. In severa! cases, especially in older maps, elevations, latitude/longitude readings, and sometimes Division 2 names were, for the same páramo, different on different maps. Therefore, it must be remembered that sorne of the entries are approximate, although I ha ve tried to keep political divisions current. I have used only proper names and not vague localities such as "shrub páramo between Loja and Zamora." When a páramo name could not be located ( on a map) or confirmed (by literature or other sources), it is simply listed without further comment and the entry is incomplete. Finally, in the "Notes" entry it was not always possible to give precise páramo localities that included reference to a well-known city or cities found on all maps; therefore, whenever possible, one or more cities are given in the entry and these, used together with the geographical coordinates, should help to pinpoint the locality in q~estion. Undoubtedly there are many more local names that I have missed, and I very much welcome additions and corrections to the gazetteer.
137
138
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
[VOL. 84
Costa Rica and Panama James L. Luteyn
The páramos of Costa Rica and Pan ama are located in the Cordillera de Talamanca, the backbone of eastern Costa Rica and adjacent western Panama. About 60 km2 of páramo occur in the massive Cerro Chirripó region (Cleef & Chaverri, 1992), with a scattered few others toward the border with Panama. There the páramo is found between 3300 and 3819 m elevation and is dominated by the dwarfbamboo Chusquea subtessellata, which may forrn a ground cover of up to 60% (Kappelle, 1990). The páramo-like vegetation along the Panamerican Highway, in the region known as Cerro de La Muerte, at about 3100 m, is manmade (fire-induced), with numerous characteristic páramo plants found in exposed boggy sites, locally called "paramillo," "ciénagas," or "tuberas." The peaks of Volcán Irazú (3432 m) and Volcán Turrialba (3339 m) are páramo-like in appearance but are not true páramos. Hunter (1959) estimated that about 1000 ha (ca. 10 km2), or 0.02% ofthe totalland area, ofCosta Rica was páramo.
In Panama, the very summit ofVolcán Barú (also lrnown as Cerro Chiriquí), at 3475 m, has páramo-like vegetation, although it is now impossible to say what the original vegetation was like since the top has been bulldozed for communication towers. True undisturbed páramo do es seem to occur in Pan ama near the border with Costa Rica in areas such as Cerro Echandi (3160 m) and Cerro Fábrega (3335 m) (fide G. Davidse collection labels). Costa Rican and Panamanian national parks that inelude páramo are Chirripó (in Costa Rica) and Vulcán Barú (in Panama), both units ofthe larger Parque Internacional de la Amistad shared by the two countries. For further information about Costa Rican and Panamanian páramo vegetation see Chaverri, unpubl. data; Chaverri et al., 1997; Cleef & Chaverri, 1992; Gómez P., 1986, 1994; Hooghiemstra et al., 1992; Horn, 1.990b; Janzen, 1983; Kappelle, 1991; Vargas Ulate & Sánchez G., unpubl. data; Weber, 1958, 1959; and Weston, 1981a, 1981b.
'D 'D
N ame
Elev. (m)
Latitude
Longitude
Division 1
Division2
Notes
Aka, Cerro
3225
09°l5'N
83°l9'W
Puntarenas/Limón
Buenos Aires/Talamanca Cord. de Talamanca
Ami, Cerro
3295
09°24'N
83°30'W
San José/Limón
Pérez Zeledón/ Talamanca Cord. de Talamanca, ENE of San Isidro del General
_Amo, Cerro
3460
09°26'N
83°29'W
San José/Limón
Pérez Zeledón/ Talamanca Cord. de Talamanca; also spelled Amuo (fide L. D. Gómez, pers. comm.)
Apri,Cerro
3190
09°l5'N
83°03'W
San José/Limón
Pérez Zeledón/ Talamanca Cord. de Talamanca, Cerro Kámuk massif
Asunción, Cerro
3396
09°35'N
83°45'W
San J osé/Cartago
Pérez Zeledón/Paraíso
Barú, Volcán
3475
08°48'N
82°32'W
Chiriquí
Boro, Cerro
3430
09°29'N
83°28'W
Limón
Talamanca
Brunka, Cerro
3030
09°2l'N
83°2l'W
Puntarenas/Limón
Buenos Aires/Talamanca Cord. de Talamanca
Buena Vista, Cerro (de)
3491
09°33'N
83°45'W
San José
Pérez Zeledon
See Cerro Amo
Amuo, Cerro
Cord. de Tal aman ca, part of Cerro Buena Vista massif, highest point along Interamerican Hwy. PANAMA, Cord. de Talamanca, NNW ofDavíd; also called Volcán Chiriquí Cord. de Talamanca Cord. de Talamanca, along Interamerican Hwy., W ofCerros Cuericí, Cerro Asunción is part ofthis massif; sometimes spelled Buenavista; also called Cerro de La Muerte, Cerro (Las) Buvis (Buvis is an abbreviation for BUena VISta), or Pico Páramo
Buvis, Cerro (Las)
See Cerro Buena Vista
Chiriquí, Volcán
See Vólcán Barú 3819
09°32'N
83°29'W
Cartago/San José/Limón Turrialba/Pérez Zeledón/ Cord. de Talamanca, SE of Cartago, NE of San Isidro, SE of Limón Cerro Cuericí, NW ofCerro Dúrika; also called Cerro Chirripó, Macizo de Chirripó Grande, or Monte Walker
3765
09°28'N
83°29'W
San José
Pérez Zeledón
3394
09°35'N
83°38'W
Cartago/San José
Paraíso/Pérez Zeledó!l
3280
09°2l'N
83 o 18'W
Limón/Puntarenas
Talamanca/Buenos Aires Cord. de Talamanca, SE ofCerro Chirripó, N ofBuenos Aires, WNW of Cerro Cruz del Obispo, E of Cerro Ena, W of Cerro Utyúm
Cord. de Talamanca, Cerro Chirripó Grandemassif, part ofFila Terbi system Cord. de Talamanca; also called Cerro Páramo; equals Cerro Chirripocillo See Pmo. Las Vueltas
Dos Amigos, Pmo. Dúrika, Cerro
~
oC/l
See Cerro Utyúm; notpáramo (fide L. D. Gómez, pers. comm.)
Cruz del Obispo, Cerro Cuericí, Cerro
~
>--
See Cerro Chirripó Grande
Chirripó, Cerro Crestón, Cerro
"'>--
See Cerro Cuericí
Chirripocillo, Cerro Chirripó Grande, Cerro
~
::;:; 'D
N ame Echandi, Cerro
Elev. (m)
Latitude
Longitude
Division 1
Division 2
Notes
3160
09°02'N
82°49'W
Puntarenas//Bocas del Toro (PA)
Coto Brus/?
COSTA RJCAIPANAMA border, Cord. de Talamanca
-
..,. o
Elí,Cerro
3097
09°23'N
83°20'W
Puntarenas/Limón
Talamanca/Buenos Aires
Ena, Cerro
3426
09°22'N
83°26'W
San José/Limón
Pérez Zeledón/ Talamanca Cord. de Talamanca, E of San Isidro del General
Estaquero, Cerro
3270
09°37'N
83°46'W
San J osé/Cartago
Fábrega, Cerro
3335
09°07'N
82°52'W
Bocas del Toro
Frío, Cerro
3471
09°33'N
83°46'W
San José
Pérez Zeledón
Cord. de Talamanca; partofBuena Vistamassif; not certain ifpáramo
Irazú, Volcán (de)
3452
09°59'N
83°53'W
Cartago
Oreamuno
Cord. Central; páramo vegetation or paramillo (fide L. D. Gómez, 1986)
S.:: tr1 S.::
Jaboncillo, Pmo.
3177
09°36'N
83 °47'W
San J osé/Cartago
Cord. de Talamanca; al so caiied Cerro J abonciiio
Kámuk, Cerro
3549
09°l6'N
83 °02'W
Limón
Ta!amanca
Cord. de Talamanca
:;:;
Lagos, Cerro
3761
Limón
Limón
Cord. de Talamanca, Cerro Chirrípó Grande massif; may equal Cerro Laguna
Laguna, Cerro
3750
09°30'N
83°29'W
Limón
Limón
Cord. de Talamanca, part ofFila Norte system
Lohmann, Cerro
3290
09°24'N
83°3l'W
San José
Pérez Zeledón
Cord. de Talamanca
ztr1
Loma Larga, Pmo. de la
3741
09°30'N
83°30'W
Limón
Limón
Cord. de Talamanca, Cerro Chirrípó massif, a fila above Valle Ancho, N ofCerro Chirrípó
o-<
P ANAMA, Cord. de Talamanca
Macizo de Chirripó Grande
See ·cerro Chirrípó Grande
Muerte, Cerro de La
See Cerro Buena Vista
Noreste, Pico
3739
Nudo, Cerro
3762
Nuevo, Cerro
3700
09°28'N
09°29'N
83°29'W
83°29'W
San José/Limón
Pérez Zeledón/Limón
Cartgo/San José
Turríalba/Pérez Zeledón Cord. Talamanca, Cerro Chirrípó massif, highest point along Fila de Urán, NW from Cerro Chirrípó
San José
Pérez Zeledón
Páramo, Cerro ( 1)
o'rj
...,
::e: tr1
~
~ tll o ..., >-z ñ >r Q
>;;o t:l
tr1
z
See El Páramo 3470
09°33'N
83°46'W
San José
Páramo, Pico Paso de los Indios
Cord. de Talamanca, Cerro Chirrípó Grande massif See Cerros Cuericí; perhaps not a real cerro just an indication of vegetation type; does not equal Cerro Cuericí (fide J. F. Morales, pers. comm.)
Páramo, Cerro (2) Páramo, El
Cord. de Talamanca
o
!Zl
Cord. de Talamanca; also cailed Cerro Páramo (2); Pittier coilecting locality; uncertain iftrue páramo See Cerro Buena Vista; perhaps not a place but an indication of vegetation type
3275
San José/Cartago
Pérez Zeledón/Turríalba Cord. de Talamanca, Cerro Chirrípó Grande massif, a long rd. used by indigenous peoples, not a particular place (fide J. Sánchez G., pers. comm.)
<o !:"'
..,. 00
N ame
Elev. (m)
Latitud e
Longitud e Division 1
Division 2
Notes
3800
09°29'N
83°29'W
San José/Limón
Pérez Zeledón/Limón
Cord. de Tal aman ca, Cerro Chirripó Grande massif; also ca11ed Pico Pirámide
Sabana de Los Leones
3250
09°25'N
83°3l'W
San José
Pérez Zeledón
Cord. de Talamanca, Cerro Chirripó Grande massif, SSW of Cerro Chirripó; also called Va11e de Los Leones or Sabana Chirripó
Sábila, Cerro
34ll
09°35'N
83°46'W
San J osé/Cartago
Sákira, Cerro
3417
09°36'N
83°46'W
Cartago
Siruska, Cerro
3408
Pirámide, Cerro Sabana Chirripó
See Sabana de Los Leones
Cord. de Talamanca Paraíso
Cord. de Talamanca, near Asunción Cord. de Talamanca, part ofBuena Vista massif; not sure if páramo
Sureste, Pico
3760
09°28'N
83°29'W
San José/Limón
Pérez Zeledón/Limón
Terbi, Cerro
3765
09°28'N
83 °30'W
San José/Limón
PérezZeledón/Talamanca Cord. de Talamanca, Cerro Chirripó Grande massif, SSW of Cerro Chirripó
Truncado, Cerro
3685
09°29'N
83°29'W
Limón
Limón
Cartago San J osé/Cartago
Turrialba, Volcán (de)
3329
10°02'N
83°46'W
Urán,Cerro
3500
09°32'N
83°33'W
Cord. de Talamanca
Cord. de Talamanca, Cerro Chirripó Grande massif Cord. Central; considered páramo by J. Sánchez G. and J. F. Morales (pers. coriun.)
Pérez Zeledón/Turrialba
Cord. de Talamanca, Cerro Chirripó Grande massif, W of Cerro Chirripó Grande; also called Pico Urán or Cerro Chirripocillo (=Cerros Cuericí); 3620 m (fide J. F. Morales, pers. comm.); not equal to Cerro Chirripocillo (fide J. F. Morales, pers.comm.)
Utyúm, Cerro
3087
09°l9'N
83°10'W
Puntarenas/Limón
Buenos Aires/Talamanca Cord. de Talamanca; J. Sánchez G. (pers. comm.) says this is páramo and maybe includes Cerro Cruz del Obispo
Va11e de Las Morenas
3460
09°30'N
83 °29'W
Cartago/Limón
Turrialba/Limón
Cord. de Talamanca, Cerro Chirripó Grande massif, NNE of Cerro Chirripó
Valle de Los Conejos
3600
09°27'N
83°28'W
San José
Pérez Zeledón
Cord. de Talamanca, Sto SW ofCerro Chirripó Grande; 3440 m (fide J. F. Morales, pers. comm.)
Valle de Los Lagos
3500
09°30'N
83°30
San José
Pérez Zeledón
Cord. de Talamanca, Cerro Chirripó Grande massif, W of Cerro Chirripó; sorne say this equals Valle de Las Morenas-not so (fide J. F. Morales, pers. comm.)
Ventisqueros, Cerro
3812
09°28'N
83°3l'W
San José
Pérez Zeledón
Cord. de Talamanca, Cerro Chirripó Grande massif, highest peak ofFila Cemeterio de la Máquina range; al so called Pico de Los Ventisqueros
Vueltas, Pmo. (de) Las
3156
09°37'N
83 °5l'W
San José
Dota
Cord. de Talamanca, SSE ofCartago, W ofCerro Buena Vista (incl. Pmo. Dos Amigos); also known as Cerro de Las Vueltas
3399
09°35'N
83°46'W
San José/Cartago
Valle de Los Leones
"O ~
~ o CZl
S::
See Sabana de Los Leones
Walker, Monte Zacatales, Cerro
"'
"' ..:S
See Cerro Chirripó Grande Al so called Cerro de la Muerte, according to Weber (1959)
:¡;:
142
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
[VOL. 84
Colombia James L. Luteyn and Mauricio R. Gavilanes A.
The Andes mountains ofColombia are essentially divided into three ranges (or cordilleras) that diverge to the north and are separated by deep valleys. The Cordillera Central runs up the center of Colombia and contains sorne of the highest snow-capped peaks in· the country with many extensive páramos. The Cordillera Oriental runs northeast-southwest and is the broadest and most massive of the three mountain ranges. It divides with a northem spur, the Sierra de · Perijá, which runs along the Colombia-Venezuela border and slowly slopes into the Caribbean, and an eastem spur, the Cordillera de Mérida, which runs northeast into westem Venezuela. The southem end ofthe Cordillera Oriental merges with the Cordillera Central in an area known as the Macizo Colombiano. The Sierra Nevada de Santa Marta is an isolated massif in northem Colombia, separated to the northwest of the Cordillera Oriental by the Magdalena-Cesar depression. It has many very high páramo regions that are poorly known (Cabot, 1939; Cleef & Rangel Ch., 1984; Rangel Ch. et al., 1982; Seifriz, 1934, 1937; Sievers, 1888; Van der Hammen & Ruiz, 1984). The Cordillera Occidental is a relatively low mountain range of recent origin. It has only a few páramos of limited size in the states of Antioquia (Páramos de Caramanta, Frontino, and Paramillo) and Valle del Cauca (the Farallones de Cali region). One small páramo in the southem part ofValle del Cauca was only recently discovered (M. Becking et al., 1997). In southem Colombia (Department of Nariño ), the three cordilleras join into a mass of high páramos
known as the Nudo de Los Pastos that extends into northem Ecuador (Páramos del Angel). The páramos of Colombia are centered in the northem half ofthe Cordillera Oriental, in the departments of Meta, Cundinamarca, Boyacá, and Santander, and in theCordillera Central, in the departments of Cauca, Valle del Cauca, and the Qüindío-Tolima-Caldas border regions. Cleef (in Cleef & Chaverri, 1992) estimated that the Cordillera Oriental alone has 100 km2 of páramo, although the overall area occupied by páramo in Colombia is unknown. Most páramos occur above 3000 m in Colombia, but the lowest páramos occur in azonal habitats such as boggy areas as low as 2380 m in the Cordillera Central (Huila: Turbera de La Candelaria, an open boggy vegetation with Espeletia; se e Rangel Ch. & Lozano C., 1986) and 2650 m in the Cordillera Occidental (Cauca: Paramillo Cerro California; see Becking et al., 1997). Colombian national parks that include páramo are Chingaza, El Cocuy, Los Farrallones de Cali, Las Hermosas, Nevado de Huila, Los Nevados, Paramillo, Pisba, Puracé, Sierra Nevada de Santa Marta, Sumapaz, and Tamá (cf. Sánchez P. & HemándezCamacho, 1995). For additional discussions ofpáramo in Colombia, see Cleef, 1981b; Cuatrecasas, 1934, 1957, 1968; Guhl, 1982; Lozano C. & Schnetter, 1976; Rangel Ch., 1991a; Sturm & Rangel Ch., 1985; and van der Hammen and associates, various works including the ECOANDES books.
::0
'D
N ame Achupallas, Pmo. (de)
Elev. (m)
Latitude
Longitude
Division 1
4050
01 o35'N
76°47'W
Nariño/Cauca
Aconta, Pmo.
Division2
Boyacá
Aquitania
Cundinamarca
D.F.
Boyacá
Tota
See Cerro Pan de Azúcar 3500
05°00'N
74°!4'W
Aguacatal, Pmo.
Cord. Oriental, NW ofBogotá See Pmo. de Herveo Cord. Oriental
Aguaná, Pmo.
3500
Agüero, Pmo. de
3950
05°59'N
73°05'W
Boyacá
Cord. Oriental, Virolín-Duitamard.; also calledPmo. de Avero; probably part of the larger Pmo. de La Rusia
Albarracín, Pmo.
3000
05°!6'N
73 °35'W
Cundinamarca/Boyacá
Cord. Oriental, Bogotá-Tunja rd., S of Pmo. Rabanal and connected with ít
00°5l'N
77°13'W
Nariño
Alcalde, Cerro
3500
Alfombras, Pmo. (Las)
4000
Alfombras, Pmo. (de Las)
3800
05°20'N
73°05'W
Almaguer, Pmo. de
S ofPasto, S ofLaguna de La Cocha
Santander
Onzaga
Cord. Oriental
Boyacá
Aquitania
Cord. Oriental, SE ofTunja, NE ofMiraflores, SE ofPmo. de Sianchoqué and connected with it
Cauca
Macizo Colombiano; Bonpland locality
Almorzadero, Pmo. del (1)
2100
06°29'N
76°25'W
Antioquia
Urrao
Cord. Occidental, Parque Nacional de las Orquídeas
Almorzadero, Pmo. (de)! (2)
4475
06°57'N
72°4l'W
Santander
Cerrito
Cord. Oriental, 8 km ESE ofBucaramanga, NE of Guaca
Alpes, Pmo. (de Los)
3100
04°20'N
75°30'W
Tolima
!bagué & Cajamarca
Cord. Central, WSW oflbagué, E ofPmo. de Don Simón and connected with it
Altar, Cerro
3620
01 °IO'N
7r36'W
Nariño
Túquerres
WSW ofPasto, N ofTúquerres; also called Pmo. Quitasol
Boyacá (?)
Belén(?)
Cord. Oriental
Nariño
Pasto & Buesaco
Cord. Centro-Oriental, NE ofPasto
Alto Belén, Pmo. de 01 °l5'N
77°!2'W
Alto Las Animas
3500
Alto Pararnillo
3350
Caldas
Riosucio
Cord. Central
Alto Tetica
3700
Boyacá
Sativanorte
Cord. Oriental
Alto Tidaqui
3500
Boyacá
Socha
Cord. Oriental
Alto Tirapa
3600
Boyacá
Chita
Cord. Oriental
Alto Torrecillas
3800
N Santander
Mutiscua
Cord. Oriental
Alto Zapayurco
3500
Nariño
Pasto
01 °l2'N
77°08'W
Alto de Canutos
"ti
>~ ;$ o
(/)
Cord. Centro-Oriental, E ofPasto See Pmo. de Canutos
Alto de Sabanazos Alto de Socaca
~
Cord. Central, ENE of La Cruz, N of Pmo. Tajumbina and connected with it, incL Cerro Petacas and Cerro Alumbra!
Agua Blanca, Cerro Agua Blanca, Pmo. de
Notes
See Pmo. de Sabanazo 3850
Magdalena
Aracataca
Sierra Nevada de Santa Marta
_¡:,.
w
Elev. (m)
Latitude
Longitude Division 1
Alto, Pmo.
3400
05°08'N
74°0l'W
Amarillo, Volcán
4500
N ame
Division 2
Notes
El Cocuy & Chita
Cord. Oriental
Cundinamarca
Cord. Oriental, Zipaquirá-Pacho rd., N ofZipaquirá
Boyacá
Amiami, Cerro
4500
Magdalena
Santa Marta
Sierra Nevada de Santa Marta
Anagá, Pmo. (de)
3700
06°53'N
72°39'W
N Santander
Cerrito
Cord. Oriental, NE ofCerrito
Andabobos, Pmo.
3700
04 °07'N
74°l5'W
Cundinamarca
San Bernardo
Cord. Oriental, W border ofPmo. de Sumapaz, ESE of Fusagasugá, incl. Alto Andabobos (4050 m)
Andes, Pmo. Los
3800
03°53'N
75°5l'W
Valle/Tolima
Angimalúa, Cerro
4000
Cord. Central, ESE of Santa Lucía, near Pmos. Rancho Largo and La Soledad
La Guajira
Riohacha
Angostura, Pmo. de
til
See map in Hettner, 1892; and Guhl, 1966: 69
o
4242
01 °30'N
76°48'W
Nariño/Cauca
LaCruz&El Tablón/Santa Rosa
Animas, Pmo. (de Las)
3700
04°05'N
74°ll'W
Cundinamarca
Gutiérrez
Cord. Oriental, part ofPmo. de Sumapaz
Apio, Pmo. El
3400
01 °56'N
76°43'W
Cauca
Vereda Florida
Macizo Colombiano, NE ofSan Sebastián, ENE of Almaguer
Cord. Central-Oriental, NE ofPasto, NE ofVolcán Doña Juana, continuation of Cerro Petacas
Arboleda, Pmo.
3350
05°30'N
75°!5'W
Antioquia
Arboledas, Pmo. de
4100
07"27'N
72°54'W
N Santander/Santander
Cord. Oriental, slopes of Cerro El Viejo, SW of Cúcuta, NE of Bucaramanga ·
Arbolito, Pmo. El
3700
02°08'N
76°36'W
Cauca
Nudo de Los Pastos, NW side ofVolcán de Sotará
Arcabuco, Pmo. de
3000
Arenales, Pmo. de
3500
07"08'N
72°55'W
Arnical, Pmo. de Arrabal, Pmo.
2380
02°!6'N
76°!5'W
Sonsón & N ariño
.
Santander
Tona
(/l
o'1:1 ..., ~
til
z
til
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Cord. Oriental, between Arcabuco and Tunja Cord. Oriental, SW ofBerlín
(")
Cord. Oriental, E ofChicamocha Valley
Huila
Cord:Central, W Volcán Purace, near Santa Leticia, Finca La Candelaria
2:;
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t:"'
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§ ztil
Locality unknown; given in the type description of Eryngium moritzii Wolff (Moritz 1146) 3500
04°20'N
73 °47'W
Cundinamarca/Meta
Quetame/El Calvario
Cord. Oriental, N ofPmo. (de) Las Burras and connected with it, SE ofBogotá, E ofCáqueza See Pmo. de Agüero
A vero, Pmo. de
See Pmo. Avión
Avión, Cerro (del) Avión, Pmo.
~
so
o ~ 1:0 ...,o >
Boyacá
Asucati, Pmo. de Atravesado, Pmo. (El)
Cord. Central, N bfPmo. SanFéliz and connected with it, W of La Dorada and Arboleda, S ofSonsón
Nariño
Asaltantes, Pmo. de Los
~
Sierra Nevada de Santa Marta
Animas, Cerro (de Las)
Boyacá
:;;: .¡,.
3550
Magdalena
Border with Venezuela, Sierra de Perijá, E ofManaure; also called Cerro (del) Avión; called Monte Plano on Venezuelan side
<o r-'
00
.¡,.
N ame
Elev. (m)
Latitud e
Longitude
Division l
Division2
Notes
Azuayo, Cerro
3600
00°50'N
77°24'W
Nariño
Córdoba
Cord. Centro-Oriental, NE ofipiales
Azufra!, Pmo.
4070
01 °04'N
77°4l'W
Nariño
Sapuyes, Mallama & Santa Cruz
W ofTúquerres, WSW ofPasto (Volcán Azufra! reaches 4070 m)
07"39'N
72°57'W
N Santander
Arboledas
Cord. Oriental, W of Arboledas; also spelled Bogueche
Bagueche, Pmo. de
3500
Balcón de Bellavista
3700
Barajas, Pmo. (de) Las
3500
04°44'N
Barbillas, Pmo. (de)
3400
Barragán, Pmo.(!) Barragán, Pmo. (2)
Magdalena
Sierra Nevada de Santa Marta, ca. 15 km NNE of San Sebastián
73°45'W
Cundinamarca
Cord. Oriental, SE of Embalse de Tominé, W of Gama, NE of Bogotá
01 °59'N
76°43'W
Cauca
San Sebastián & Almaguer (La Vega?)
Macizo Colombiano, NNE of Almaguer, between Río Guachicono and Valle de Las Papas
3800
04°05'N
75°45'W
Valle/Tolima
Sevilla, Tuluá & Buga/Roncesvalles
Cord. Central, NNW ofChaparral, NE ofPmo. de Miraflores and connected with it,just N ofParque Nacional Natural Las Hermosas
3750
03 °57'N
75°55'W
Valle
Cord. Central, SW ofBarragán, NE ofSanta Lucía, S ofPmo. Santa Lucía
Bavaya, Pmo. de
3550
Valle
Cord. Central, W slopes, el ose to Bugalagrande
Belén, Pmo. de
3950
Boyacá
Cord. Oriental, N ofBelén, incl. Alto de las Cruces
Bellones, Pmo. (de)
3100
02°05'N
76°39'W
Cauca
La Vega
Cord. Central
Belmira, Pmo. de
3100
06°35'N
75°32'W
Antioquia
Belmira
Cord. Central, 6-8 km from Belmira; incl. Alto de Sabanazos (or El Morro); sub páramo dominated by Espeletia and Puya
Beltrán, Pmo. de
2970
04°27'N
73°54'W
Cundinamarca
Berlín, Pmo. de
3800
07"ll'N
72°53'W
Santander
Betulia, Pmo. (de)
3900
03 °22'N
76°05'W
Valle
Bijagual, Pmo. (de)
3500
05°28'N
73°J]'W
Boyacá
Bijoacales, Pmo.
3600
04°!5'N
74°ll'W
Cundinamarca
Blanco, Pmo. (J)
3800
02°08'N
76°32'W
"O
Cord. Oriental, 27 km SE ofBogotá, E and above Fómeque; not true páramo Tona
~
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(/]
Cord. Oriental, between Pamplona and Bucaramanga, S ofBerlín Cord. Central, E ofCali, E ofPradera, S ofPmo. de Tinajas, N ofPico Horizonte
Tota & Pesca
Cord. Oriental, E of Tunja, W of Laguna de Tota; also called Pmo. Vijagual
Cauca
Sotará & Puracé
Macizo Colombiano, S ofPopayán, slopes ofCerro Canelo
Santander/Boyacá
Onzaga!?
Cord. Oriental, S ofBogotá, justE ofLaguna Chisacá
Blanco, Pmo. (2)
See Pmo. Mantilla
Bobos, Laguna Los
3800
Boca Grande, Pmo. de
3710
Boca del Mundo Nuevo, Pmo. de La Bocatoma, Valle de
ce
'-D
.'S
4100
04°l9'N
74°06'W
Cord. Oriental
Cundinamarca
Cord. Oriental, small páramo and severa! lagoons at N edge of Pmo. de Sumapaz, 35 km S ofBogotá; also spelled Bocagrande
Nariño
On Volcán Cumbal
Arauca
Cord. Oriental, Sierra Nevada del Cocuy, W slope area of páramos
:¡;: V!
Elev. (m)
Latitude
Longitude
Division 1
Bogueche, Pmo.
3000
or3s'N
72°58'W
N Santander
Cord. Oriental, NNEofBucaramanga, SW ofCúcuta, S ofPmo. de Castro and connected with it
Boquerón, Pmo. El (de)
3500
Cundinamarca
Cord. Oriental, S ofBogotá; Pmo. de Boquerón was the pass between Bogotá and Chipaque; name was used mainly before 1917
N ame
Division 2
Antioquia
Notes
Boquerón, Pmo. de
3100
Bordoncillo, Cerro El
3700
0! !4'N
7r08'W
Putumayo/Nariño
Colón/Pasto & Buesaco
20 km E ofPasto, extreme NW ofPmo. del Bordoncillo; also called Volcán Bordoncillo and Volcán Pataseos
Bordoncillo, Pmo. de(l)
3600
01 o 13'N
7r08'W
Nariño/Putumayo
Pasto/Santiago
Slopes ofVolcán Bordoncillo, 20 km E ofPasto
0
See Cerro El Bordoncillo
Bravo, Cerro
Cord. Central (?) 3800
03°28'N
76°00'W
Tolima/Valle
Cord. Central, E ofCali, ENE ofPradera, NE ofPmo. de Tinajas
Brujo, Pmo. del
Cord. Central, on Nevado del Huila
Buena Vista, Pmo. (de)
3600
Buey, Alto de
3600
Buey, Pmo. del (2)
Cau ca 75°34'W
Antioquia
4200
01 °58'N
76°37'W
Cauca/Huila
3800
or4l'N
72°57'W
Buey, Pmo. del (3) Bueyes, Pmo. Los
. Cord. Central, in Cord. Buena Vista
Valle 05°44'N
Buey, Pmo. del (l)
~
Cord. Central, subpáramo vegetation at top
Bordoncillo, Volcán Brillante, Pmo. El
...
Cord. Occidental, Los Farallones de Cali Sotará, Puracé/San Agustín
Macizo Colombiano, Parque Nacional Natural del Puracé, N of Pmo. de las Papas and connected with it, S ofPopayán
Nariño
25 km E ofPasto
N Santander
Cord. Oriental, SW of Salazar
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ztn
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Buitre, Pmo.
3800
Santander
Cerrito
Cord. Oriental
Buncuabí, Cerro
4000
La Guajira
Riohacha
Sierra Nevada de Santa Marta
Burgos, Pmo. Los
3400
Nariño
Ospina
Nudo de Los Pastos
op.
Burinchucua, Cerro
4400
Cesar
Valledupar
Sierra Nevada de Santa Marta
E3
Burras, Pmo. (de) Las
3500
04°23'N
73°48'W
Cundinamarca/Meta
Quetame/El Calvario
Cord. Oriental, S ofPmo. Atravesado and connected with it, E of Cáqueza; al so called Pmo. Los Burros
3600
01 °55'N
77°l8'W
Nariño
Punes
Cord. Centro-Oriental, SW ofPasto
Burros, Pmo. Los Caballo Rucio, Cerro
t-
ti1
z
Se e Pmo. (de) Las Burras
Cabeicaca, Cerro
4050
Aracataca
Sierra Nevada de Santa Marta
Cabrera, Cerro
3512
Ol 13'N
77°10l'W Nariño
Magdalena
Pasto & Buesaco
NE ofPasto
Cabrera, Pmo. La
3000
orl9'N
72°29'W
N Santander
Herrán & Toledo
Cord. Oriental
Cáchira, Pmo. de
3500
or50'N
73°00'W
N Santander
0
Cord. Oriental, N ofPmo. de Guerrero and connected with it; 60 km W ofCúcuta, 10 km S ofVillacaro, N ofCáchira
<o ~
... 00
N ame
Elev. (m)
Latitude
Longitud e
Division 1
Division2
Notes
Cachiri, Pmo. de
3-4000
07"28'N
73°0l'W
Santander
Cord. Oriental, vicinity ofCerro El Viejo, ca. 45 km N ofBucaramanga and abo ve village ofCachirí; coordinates approximate
Cadillal, Pmo. de
3800
05°55'N
72°38'W
Boyacá
Cord. Oriental, SSE ofSocha Viejo
Cajón, Pmo. El
3800
03°58'N
74o08'W
Meta
Cord. Oriental, E oflarger Pmo. de Sumapaz
Quin dí o
Cord. Central
Calarcá, Pmo. Calera, Pmo. de La
3000
Cundinamarca
California, Cerro
4100
Cau ca
Argelia
Cord. Oriental
California, Paramillo Cerro
3250
Cauca
Balboa/Argelia
Cord. Occidental, along Balboa-Argelia rd. at the crest ofthe cordíllera; páramo descends to 2650 m; southernmost páramo in Cord. Occidental
Calocribe, Pmos. de Laguna de
3700
Campanario, Cerro El
3700
Caña, Pmo. La
3400
02°IO'N
77"15'W
\0 \0
.::S
Cord. Oriental
Magdalena
Sierra Nevada de Santa Marta, E of Meo llaca Cord. Central, La Línea
06°06'N
72°28'W
Candelaria, Pmo. de La
Boyacá
Chita
Cord. Oriental, S ofChita Cord. Central
Huila
La Plata
Candor, Pico
4200
Boyacá
El Cocuy & Chita
Cord. Oriental
'ti
Canela, Cerro
4100
Cauca
Sotará & Puracé
Cord. Central
Canoas, Pmo.
4000
Boyacá
Chita
Cord. Oriental
S::;;:::
Cañutal, Pmo. del Alto del
3400
Boyacá
Canutos, Pmo. de
3700
06°04'N
72°55'W
Capilla, Pmo. Capuchino, Pmo. (de) El
3400
Caracas, Cerro
3900
01 °57'N
73°34'W
Cord. Oriental, between Soatá and Cocuy; also called Alto del Cañutal
Boyacá Socha
Cord. Oriental
Putumayo
Santiago
El Encano-Sibundoy rd.
La Guajira
Sierra Nevada de Santa Marta, N slope, SE ofSan Miguel; also called Cuchilla Caracas
Caramanta, Cerro
See Pmo. Caramanta See Pmo. Caramanta 3900
Carbonera, Cerro La
4000
05°30'N
76°00'W
Carcasí, Pmo. de
Risaralda!Antioquia! Chocó
Mistrató/Andes/Bagadó
Boyacá
Chita
Cundinamarca
Carbonera, Pmo. de La 3800
06°45'N
72°30'W
Santander
o
(/l
Cord. Oriental, W ofPmo. de Guantiva; also called Alto de Canutos
Boyacá
Caramanta, Cordillera Caramanta, Pmo.
>-
Cord. Occidental, Farallones de Citari, SW ofJardín; also caiied Cordiiiera or Cerro Caramanta Cord. Oriental See map in Hettner, 1892; and Guhl, 1966: 100; possibly the same as Pmo. de Guasca
Carcasí & Concepción
Cord. Oriental, N ofSierra Nevada del Cocuy, E ofMálaga, NE ofCarcasí
.¡:,..
-.¡
N ame
Elev. (m)
Latitud e
Longitude
Division 1
Cardenillo y Ventura, Pmo. de
4400
06°30'N
72°2Ü'W
Boyacá
Cardón, Pmo. del
3590
Division 2
Notes
:¡;: 00
Cord. Oriental, SW ofNevado de Güicán
Santander
Cord .. Oriental, close to Casanare
Carmen, Pmo. del
See Pmo. Monte de Leona Santander
Onzaga
Cord. Oriental
Carnicerías de Sianogo, Pmo.
3800
Carrizales, Pmo.
3800
03 °56'N
75°39'W
Tolima
Cascabel, Pmo. de
3000
01 °40'N
76°34'W
Cau ca
Santa Rosa
Macizo Colombiano
Cascabelito, Pmo. de
3300
01 °40'N
76°34'W
Cauca
Santa Rosa
Macizo Colombiano
Casitas, Lagunilla de Las
3700
Cau ca
Casitas, Pmo. del Alto de Las
3800
Cau ca
Castillo, Pico El
5400
Boyacá
Güicán
Cord. Oriental, Sierra Nevada del Cocuy
Castro, Pmo. de
3600
Or4l'N
72°55'W
N Santander
Salazar & Arboledas
Cord. Oriental, S ofPmo. de Pringador and connected with it, WSW ofCúcuta, NW of Arboladas
Catanga, Pmo.
3000
0! 51'N
76°30'W
Cauca/Huila
Cazadero, Pmo.
3500
06°!0'N
72°42'W
Boyacá
Cedral, Pmo. El
3500
04°08'N
74°07'W
Cundinamarca
Centellón, Pmo. El
3600
Centro, Pmo.
3100
0
Cord. Central, SSW ofRoncesvalles
Cord. Central, between Quebradas de San Paulino and del López Cord. Central, W slopes, near del Río Palo
Macizo Colombiano, SE ofPmo. de las Papas and connected with it, N ofSanta Rosa · Cord. Oriental Gutiérrez
Cord. Oriental, S ofBogotá, SE ofFusagasugá, N ofPmo. de Sumapaz
Boyacá (?) 05°29'N
73°29'W
Cord. Oriental, SSW ofTunja, SE ofSamacá; village, probably notpáramo See Pmo. de Guantiva
Cerinza, Pmo. de 3500
Cundinamarca
Chaquiro, Pmo. de
3200
Bolívar/Antioquia
Charambud, Cerro
3400
00°58'N
77°36'W
Nariño
Chasques, Pmo. (de, Los)
3500
05° 13'N
73 °3l'W
Cundinamarca
Chía, Pmo. de
3600
06°59'N
72°38'W
Santander
Chicó, Pmo. del
3300
Chiles, Volcán
4748
00°48'N
7rss·w
Chilí, Pmo. (de)
3930
04°l5'N
75°38'W
o
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(/)
o71
...,
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~
to
Boyacá
Chamizale, Pmo.
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Fómeque
Pupiales
Cord. Oriental, 3.5 km SW ofLaguna de Chingaza
o..., > 2:1 ()
r>
Q
Cord. Occidental, Urrao-Antisales rd., probably currently the same as is called Paramillo (Antioquia)
>
Cord. Occidental
z
~
trl
Cord. Oriental, above Villapinzón, fuentes del Río Bogotá Cerrito
Cord. Oriental
Nariño (CO)/Carchi (EC)
Cumbal/Carchi
Nudo de Los Pastos, W of Ipiales; along Colombia/Ecuador border; also called Nevado del Chiles
Quindío/Tolima
Pijao/Roncesvalles
Cord. Central, N ofPmo. de Cumbarco and connected with it, E ofSevilla, SE ofCaicedonia
Cundinamarca
Cord. Oriental, Macizo de Bogotá
<o ~
00
.¡:,.
N ame
Elev. (m)
Latitud e
Longitude
Division 1
Division 2
Notes
Nariño
Aldana, Sapuyes & Pupia1es
Cord. Occidental, Nudo de Los Pastos
73°33'W
La Guajira
Riohacha
Sierra Nevada de Santa Marta; not páramo
76°55'W
Nariño
San Bernardo
Cord. Centro-Oriental, N of San Bernardo
01 °08'N
77°13'W
Nariño
Pasto
SE of Pasto, W of Laguna de La Cocha, N of Pmo. de Guapuscal and connected with it; also called Pmo. de Siquitán or Serranía del Tábano
3800
03°24'N
76°00'W
Valle/Tolima
Palmira/Rioblanco
Cord. Central, N ofPmo. de Las Hermosas and connected with it, E ofPalmira
Chingaza, Pmo. (de)
3500
04°31'N
73°45'W
Cundinamarca
Fómeque
Cord. Oriental, NE of Choachí, S of Pmo. de las Barajas and connected with it, 25 km E ofBogotá; also spelled Chinguasa or Chingasa in early days
Chipaque, Pmo. de
3300
Chirigua, Pmo. de
4000
l0°56'N
73°21'W
La Guajira
Chirugua, Pmo. de
4570
10°51'N
73°41'W
Magdalena
3450
00°57'N
77"39'W
Chimalongo, Pmo.
2500
Chimayoy, Cerro
3700
l1°0l'N 01 o29'N
Chimbalán, Pmo.
3500
Chinche, Pmo. de
Chiltazón, Pmo. (de)
'D 'D
~
Cord. Oriental
Cundinamarca Riohacha
Sierra Nevada de Santa Marta, N slope; also called Alto de Chírua and Pmo. de Chirucua; páramo starts about 3000 m Sierra Nevada de Santa Marta, at headwaters ofMacotama River; probab1y the same as Pmo. de Chirigua; also spelled Chiruqua in Todd & Carriker, 1922 See Pmo. de Chisacá
Chisacal, Pmo. Chisacá, Pmo. de
4120
Cundinamarca
Cord. Oriental, also called Pmo. Chisacal, spur ofPmo. de Sumapaz, 45 km SW ofBogotá on rd. to Usme
Chitá, Pmo. de
3700
Boyacá
Cord. Oriental, Sierra Nevada del Cocuy, headwater ofRío Casanare; Nevado goes to 5400 m
Chitagá, Pmo. de
3600
07"11'N
72°36'W
N Santander
Chitagá
Cord. Oriental, NE ofChitagá
Choachí, Pmo.
3500
04°33'N
73°58'W
Cundinamarca
Choachí, Ubaque & Bogotá
Cord. Oriental, E ofPmo. Cruz Verde andconnected withit, 12 km SE ofBogotá
Chocontá, Pmo. de
2830
Chontales, Pmo. de
4000
05°50'N
-o
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o
(/)
Cord. Oriental; probably not a Páramo
Cundinamarca 73°10'W
Santander/Boyacá
Charalá & Encino/Duitama& Paipa
Cord. Oriental, SW ofPmo. de la Rusia and connected with it, E ofMoniquirá
Chorrera, Cerro de La
3400
00°52'N
77°22'W
Nariño
Córdoba
Cord. Centro-Oriental, E oflpiales
Chunchullo, Pmo. del
3000
01 o43'N
76°45'W
Cau ca
San Sabastián, Santa Rosa & Bolívar
Cord. de Los Andes; also spelled Chunchulo; also called Pmo. de Granadillo
Chusca!, Pmo. El
3850
05°50'N.
72°45'W
Boyacá
Tasco
Cord. Oriental, NE ofGameza, SE ofTasco
Chusque, Pmo. El
4200
06°20'N
72°20'W
Arauca/Boyacá
Cord. Oriental, SW comer ofParq. Nac. Nat. El Cocuy, SE of El Cocuy
:¡;: 'D
Elev. (m)
Latitud e
Longitude
Division 1
Cinera, Pmo. de
3400
07"32'N
72°55'W
N Santander
Cisne, Nevado del
5200
04°49'N
75°23'W
Caldas
Citará, Pmo. de
3900
05°45'N
76°05'W
Antioquia!Chocó
Cord. Occidental, Farallones de Citará (also called Farallones de Chocó), W of Andes, incl. Cerro San Nicolás, Cerro San Fernando, and Cerro Caramanta (the highest peak in the Farallones is at ca. 3900 m)
Clarín, Pmo. El
3500
04°02'N
74°05'W
Meta/Cundinamarca
Cord. Oriental, NE of the larger Pmo. de Sumapaz
Coatí, Pmo. de
2870
Boyacá
Cord. Oriental, Tl!I\ia
Cobardes, Pmo. de Los
3500
06°35'N
73°27'W
Santander
Cord. Oriental, SSW ofBucaramanga, W ofBarichara
Cobre, Pmo. del
3613
07"19'N
72°29'W
N Santander
Toledo
Cord. Oriental, within Parque Nacional Natural Tamá
Coconucos, Pmo. de
4500
02°16'N
76°22'W
Cauca!Huila
Puracé/La Plata
Cord. Central, SE of Popayán, part of Sierra Nevada de Los Coconucos (incl. Volcán Puracé, V. Coconucos, Pico de Paletará, Cerro Pan de Azúcar, Cerro Pelado)
N ame
Cocuy, Sierra Nevada del
5493
06°25'N
72°18'W
Boyacá
Division2
Notes
Villamaría
Güicán, Cocuy & Panqueba
Cord. Central, S ofNevado del Ruiz
Cord. Oriental, also called Sierra Nevada de Chita; highest point is Nevado del Cocuy See Sierra Nevada del Cocuy
10°5l'N
73°35'W
Magdalena
3400
01 °00'N
77°45'W
Nariño
Collagá, Pmo. de
3800
07°05'N
72°54'W
N Santander
Cord. Oriental, N of Santa Bárbara, SW of Berlín
Colombiano, Macizo
3500
01 °30'N
76°30'W
Cauca!Huila
Also called Gran Macizo Colombiano; S ofPopayán, connects and incl. Pmo. de Las Papas, Pmo. Catanga, Cerro Cusiyaco, and Alto de Puinchano
Coloradas, Pmo. de Las
3900
Santander
Cord. Oriental, above La Baja
Colorado, Pmo. de
3800
Colorados, Pmo. Los(!) Colorados, Pmo. Los (2) Cóncavo, Pmo. (El)
4400
06°23'N
4350
06°08'N
5150
Cogua, Pmo. de
3650
Colimba, Cerro de
s;: m s;: o ~ CJ:l
o
'Tj
::e: ""'
m
zm ~ >-<
Sierra Nev<;ida de Santa Marta, NW ofValledupar
o
Cord. Occidental, N extension ofPmo. de Cumbal, ca. 10 km W ofTúquerres
o
~
07"04'N
72°47'W
Guachucal
N Santander
Silos
Cord. Oriental, NNE ofBaraya
3600
Boyacá
Monguí
Cord. Oriental
3400
Boyacá
Susacón
Cord. Oriental
72°20'W
Boyacá
Güicán & El Cocuy
Cord. Oriental, Sierra Nevada del Cocuy, W of Laguna Grande (de los Verdes), NE ofMorro El Negro
Santander
Cord. Oriental
72°53'W
Boyacá
Cord. Oriental, Cord. de Consuelo, SW ofCordillera Guantivá and connected with it, NE ofCordillera Barrosas and connected with it, 20 km NE ofBelén
Concepción, Pmo. de Consuelo, Pmo. del
o
Cord. Oriental, W ofCucutilla
Cocuy, Nevado del Codazzi, Pico
V.
tJ:j
""' z>
ñ
> r o > (;§
zm
<o r'
00 ~
Name
Elev. (m)
Latitud e
Longitude
Division 1
Division2
Contento, Pmo. de
Notes See Pmo. de Méndez
Coper, Pmo. de
Boyacá
Coper
Cord. Oriental, W ofTunja
3600
05°19'N
73 °02'W
Boyacá
Aquitania, Zetaquirá, San Eduardo & Tota
Cord. Oriental, SW ofLaguna de Tota, SE ofTunja
Corralitos, Pmo.
4600
06°26'N
72°20'W
Boyacá
Cortadera, Pmo. de La (1)
3800
05°32'N
73 °07'W
Boyacá
Cortadera, Pmo. de La (2)
3400
Crisol, Pmo. El
3200
Cristal, Pmo. El
3550
Cristóbal Colón, Pico
5775
Magdalena
Santa Marta & Aracataca Sierra Nevada de Santa Marta, NW ofValledupar
Cruces, Alto de Las (1)
3900
Boyacá
Gámeza
Cord. Oriental
Cruces, Alto de Las (2)
3850
Boyacá
Socha
Cord. Oriental
Cruces, Pmo. (de) Las
4000
Cruces, Pmo. Alto de las
3380
Cruz Colorada, Pmo. de
3700
06°25'N
72°35'W
Santander/Boyacá
Onzaga, Soatá/Susacón
Cord. Oriental, NE ofCordillera Guantivá and connected with it, ESE of Socorro
' 3500
04°34'N
74°02'W
Cundinamarca
Ubaque, Choachí & Bogotá
Cord. Oriental, ca. 10 km ESE ofBogotá, W ofPmo. Choachí and connected with it, SW of Tinjacá; probably the same as Pmo. El Ve¡jón
Coroneles, Pmo.
Cruz Verde, Pmo. (de)
Cord. Oriental, ESE ofGüicán, W ofNevado El Cóncavo Toca
Nariño 05°45'N
72°35'W
Boyacá
05°32'N
73°42'W
73°08'W
Cruz, Pmo. de La
Boyacá
Cord. Oriental, NE ofPmo. de Sianchoque and connected with it, S ofPmo. de Tibamá and connected with it, E ofTunja
Cundinamarca
Cord. Oriental, Macizo de Bogotá, between La Viga and Diego Largo
N Santander
""' > ~ 3::: o (/)
Cord. Oriental, near Ocaña See Pmo. de Cruz Verde
3400
Cuchero, Pmo. de Cuchijao, Pmo.
Cord. Oriental Cord. Oriental, 2 km W ofPmo. de Chingaza, S ofLaguna del Ami cal
Cruzverde, Pmo. de Cuaspud, Cerro
Cord. Oriental, W ofLaguna de Tota, E ofTunja Km 23-23 on lpiales-La Victoria rd.
Pisba
Cundinamarca 10°5l'N
'O 'O 'O
~
00°5l'N
7r43'W
Nariño
Cario sama
Santander!N Santander (?)
Cord. Occidental, W oflpiales Cord. Oriental
3900
Boyacá
Monguí, Mongua, Aqui- Cord. Oriental tania & Labranzagrande
Cueva, Pmo. Alto de la
4030
Boyacá
El Cocuy & Güicán
Cord. Oriental, Sierra Nevada del Cocuy
Cuigueca, Pico
4100
Magdalena
Aracataca
Sierra Nevada de Santa Marta
Cuchilla Caracas
See Cerro Caracas
::;;
Elev. (m)
Latitude
Longitude Division 1
Division2
Notes
Cumbal, Pmo. de
3500+
00°57'N
1rs2w
Nariño
Cumbal, Guachucal & Mallama
Nudo de Los Pastos, slopes ofNevado (de) El Cumbal (4764 m), 18 km NE ofVolcán de Chiles
Cumbarco, Pmo. de
3700
04 °07'N
75°42'W
Tolima!Quindío
Roncesvalles/Génova
Cord. Central, N ofPmo. Barragán and connected with it, SE of Sevilla, SW oflbagué
Curiba, Pmo. (de)
4100
10°5l'N
73°3l'W
Cesar
Valledupar
Sierra Nevada de Santa Marta; also called Pmo. de Curigua; incl. Laguna Curigua; also spelled Curibia
Curies, Pmo. de Los
3500
05a32'N
72°52'W
Boyacá
Aquitania
Cord. Oriental, E ofLaguna de Tota
Cusirí, Cerro
4750
06°2l'N
72°2l'W
Boyacá!Arauca
Cusiyaco, Cerro
4000
Cutanga, Pmo. de
3600
Dagua, Pmo.
3800
Dantas, Pmo. Las
3300
N ame
See Pmo. de Curiba
Curigua, Pmo. de
01 °50'N
04 a07'N
76°28'W
74a04'W
Dedo de Dios, Cerro Delicias, Pmo. de Las
3800
02°38'N
76°l4'W
Cord. Oriental, Sierra Nevada del Cocuy
Cauca!Huila
Santa Rosa & San Sebastián/San Agustín
Macizo Colombiano
Cauca!Huila
Santa Rosa/San Agustín
Macizo Colombiano, S ofPopayán, W ofSan Agustín, S tip of ParqueNaciona!Natura!Puracé;partofVolcánCutanga(4300m)
Boyacá
Monguí
Cord. Oriental
Cundinamarca
Diamante, Pmo.
3850
. Cord. Oriental, W ofVillavicencio, N ofPmo. La Leona
75°50'W
ztn
::E
Cau ca
Inzá & Silvia
Cord. Ceñtral;.S ofPmo. de Santo Domingo and connected with it; E ofSilvia; NE ofPopayán
::0
Close to Los Cudos; also called Pmo. Delirio Susacón
Cord. Oriental Cord. Central, ESE of Santa Lucía, near Pmos. Rancho Largo and La Soledad
Tolima!Valle
Diego Largo, Cerro
3540
Cundinamarca
Cord. Oriental, Macizo de Bogotá
Dirincuve Sabana, Pmo.
3250
Magdalena
Sierra Nevada de Santa Marta, souce ofthe Río Donachuí
Dolores, Pmo.
Cord. Oriental, above Barbacoas
Domínguez, Pmo. de Las
3500
03a43'N
76°07'W
Cau ca
Cord. Central, NE of Palmira, incl. Cerro Pan de Azúcar and Laguna de Las Domínguez
Domínguez, Pmo. de Los
3900
03°42'N
76a06'W
Valle
Cord. Central, NNE ofPalmira, S ofPmo. Pan de Azucar
Cundinamarca
Don Benito, Pmo. de
-l
;:¡:: tn
Cerro in the Pmo. de Las Papas
Boyacá 03°54'N
o ;;:; o"Tl (/)
See Pmo. Delicio 3300
2: tn 2:
San Agustín
Delirio, Pmo. Desaguadero, Pmo. del
N
Huila
Cauca
Delicio, Pmo.
V.
o--< r;
ttJ
o-l
;J>
z
ñ ;J> r
Q ;J>
E3 tn
z
Cord. Oriental
Don Simón, Pmo. de
3869
04°40'N
75°35'W
Quindío/Tolima
Córdoba!Cajamarca
Cord. Central, N ofPmo. de Chilí, S of Armenia, W of Caicedonia
Doña Juana, Volcán
4250
01 °30'N
76°53'W
Nariño
El Tablón & La Cruz
Cord. de Los Andes, NE of Pasto, SW of Cerro Petacas, NE part ofPmo. de Tajumbina
<o r-'
00
_¡,.
N ame
Elev. (m)
Latitud e
Longitude
Division 1
Division2
Cau ca
::0 \0 .::S Cord. Central, W slope, headwaters o fRío Palo, a stream off of
Notes
Duende, Alto del
3350
Duriameina, Pmo. de
3300
10°39'N
73°37'W
Magdalena
3250
04°45'N
74°00'W
Cundinamarca
Bogotá
Cord. Oriental, NE of Cerro Monserrate, near Bogotá
Encino, Cerro
3600
00°53'N
7r29'W
Nariño
Puerres
Cord. Centro-Oriental, E oflpiales
Eras, Pmo. (de Las)
4600
02°42'N
76°!2'W
Cauca
Silvia, Páez & Inzá
Cord. Central, el picacho Bujío is the point of 4600 m
Escoba!, Pmo. de
4200
Boyacá
La Uvita & El Cocuy
Cord. Oriental, between Soatá and Cocuy; also called Pmo. del Alto del Escoba!
Nariño
Sapuyes & Ospina
Nudo de Los Pastos
Río López . El Granizo, Pmo.
Escubilla, Pmo. La
3400
Esperanza, Pmo. La
3800
Estero, Pmo. de Los
2800
Estrella, Pmo. de La
3800
Felino, Cerro (El)
3500
03 °57'N
03°4l'N
75°49'W
75°57'W
3000
Fraile, Pmo. del
2800
or2I'N
72°39'W
Frailejón, El Pmo. del
Nariño
Macizo Colombiano, beside Pmo. de Santa Lucía, extreme S side ofLa Cocha; also called Pmo. del Estero and Pmo. de Los Esteros
Valle/Tolima
Cord. Central, SW ofPmo. de Las Hermosas and connected with it San Agustín
Part ofthe Pmo. de Las Papas
N Santander
Pamplona
Cord. Oriental, W ofPmo. de Tierra Negra and connected with it, ridge between Pamplona and Cúcuta
Putumayo
Sibundoy
Km. 82 along the Pasto-Mocoa rd.
Nariño
Between Pasto and Túquerres N ofVolcán Sotará, NE ofPmo. El Arbolito
3400
02°l2'N
76°35'W
Cauca
3600
00°45'N
77°28'W
Nariño
Potosí
Cord. Centro-Oriental, E of Ipiales
Boyacá
Labranzagrande
Cord. Oriental, Serranía el Bizcocho
Frailejonale, Pmo. de Francés, Cerro
Cord. Central, E ofSanta Lucía, SE ofBarragán, N ofPmo. La Soledad
Valle
Fontibón, Pmo. (de)
Frailejón, Pmo.
Valle/Tolima
Huila
Florida, Pmo. de
Sierra Nevada de Santa Marta, ca. 11.5 km NNE of San Sebastián
Santander
Franco, Pmo.
Cord. Oriental, near Las Vetas
Fresno, Pmo. de
Cord. Central, near Manizales
Frío, Pmo. del
Cord. Oriental, Cucutá area
Frontino, Pmo. (de) (1)
4080
Frontino, Pmo. (2)
3600
Quin dí o
Gachalá, Pmo. de
2600
Gachaneque, Pmo. (de)
3000
06°28'N
05°27'N
76°06'W
73°33'W
Antioquia
> "" ~ S:: orz¡
Urrao, Abriaquí & Caicedo
Cord. Occidental, 20 km WSW of Antioquia, above Llano Grande; incl. Picos de San José, La Horqueta, Ocaidó, Nicasio, Santana, and Alto Junco
Cundinamarca
Gacha! á
Also called Monte Batatas (Triana collection); doubtfully páramo
Boyacá
Tunja & Sáchica
Cord. Oriental, NE ofPmo. Rabanal and connected with it, SW ofPmo. de Runta and connected with it, E ofFúquene
Cord. Central, Quindío-Salento, above Guayaquil
V> (>.)
N ame
Elev. (m)
Latitude
Longitude
4200
01 °l2'N
77°20'W
06°l2'N
71 °23'W
Galeras, Pmo. del Galeras, Volcán (de)
Division2
Notes
Nariño
Pasto, Tangua & Yacuanquer
Nudo de Los Pastos, WNW ofPasto; also called Volcán Pastos and Volcán de Pasto
Canea
Totoró
Cord. Central
Boyacá
Labranzagrande
Cord. Oriental, SE ofChita, W ofLa Salina
Division 1 Nariño
Gallinazo, Pmo.
3000
Gallo, Pmo. (El) (1)
3400
V.
.p.
On Volcán Galeras, outside ofPasto
Gallo, Pmo. (El) (2)
3800
04°0l'N
74°08'W
Cundinamarca/Meta
Gutiérrez/Guamal
Cord. Oriental, S ofPmo. El Cedral and connected with it
Golondrinas, Cerro
3400
00°58'W
78°04'W
Nariño
Cumbal
Cord. Occidental, Nudo de Los Pastos, on Pmo. de Cumbal, along rd. to San Martín
Gómez, Pmo. (de) Los
3500
04°l5'N
75°30'W
Tolima
Rovira & !bagué
Cord. Central, NE ofPmo. de Chilí and connected with it, SW ofibagué
Gorgonia, Pmo. de La
3600
Nariño
Gualmatán
Cord. Occidental, NE oflpiales
[/J
Gorgua, Pmo. de
3600
Cundinamarca
Cord. Oriental, near to the Boquerón de Guargua, near to the Estación Transmisora de Televisión, above the Represa del Nuesa; also spelled Guargua
o ..., ::r: m
Gramotales, Pmo. de
3000
03°42'N
74°12'W
Meta
Cord. Oriental
Grande, Cerro
3672
05°08'N
74°0l'W
Cundinamarca
Gregoria, Pmo.
3600
04°00'N
75°48'W
Valle/Tolima
Griere, Cerro
4600
See Pmo. de Chontales and Pmo. de Chunchullo
Granadillo, Pmo. de
La Guajira
Guachaneque, Pmo. de
Tausa
Cord. Central, ENE ofSanta Lucía, SE ofBarragán, N ofPmo. La Esperanza Riohacha
Cundinamarca
Gualcalá, Cerro 01 °09'N
77°47'W
Nariño
Mallama & Santa Cruz
Nariño
Sapuyes
Gualcalla, Cerro
3950
Gualmatán, Pmo. de
3580
Guamués, Pmo. de
3400
00°50'N
77°20'W
Nariño
Guanacas, Pmo. de (Las)
3700
02°30'N
76°15'W
Cauca/Huila
Totoró/La Plata
Guanacos, Pmo. de
Sierra Nevada de Santa Marta
o
~
"rj
zm
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o ~ ttl ...,o ;¡,.
Cord. Oriental, NNE ofBogotá, N ofPmo. de Chingaza
z
See Cerro Gualcalla
l'
Cord. Occidental, NW ofTúquerres, 18 km SE ofPiedrancha, NE ofMallama, SW ofSamaniego; also called Cerro Guacalá
ñ ;¡,. Cl ;¡,.
§ m z
Cord. Central, E ofPopayán; also called Alto de Guanacas and Pmo. de Guanacos See Pmo. de Guanacas See Pmo. (de) Guerrero (l)
Guandoque, Pmo. Guantiva, Pmo. (de)
Cord. Oriént;d, E ofPmo. de Guerrero, N of Zipaquirá
3:: m 3::
4326
06ol2'N
72°45'W
Boyacá/Santander
Susacón/Onzaga
Cord. Oriental, NE ofTunja, N ofDuitama, Belén-Soatá rd., N of Pmo. La Rusia and connected with it, SW ofPmo. Cruz Colorada and connected with it; incl. Pmo. Pan de Azúcar, Pmo. de La Rusia, and Alto de La Mata; forrnerly called Pmo. de Cerinza
<o r
00
.p.
Elev. (m)
Latitud e
Longitude
Division 1
Division 2
Notes
Guapuscal, Pmo. de
3500
00°54'N
77°16'W
Nariño
Punes
NE oflpiales, S ofPasto, NE ofPmo. Las Juntas and connected with it, S ofPmo. Chimbalán and connected with it
Guardián, Pico El
5300
l0°47'N
73°38'W
Magdalena
Aracataca & Valledupar
Sierra Nevada de Santa Marta, NW ofValledupar
Guasca, Pmo. de
3400
04°54'N
73 °47'W
Cundinamarca/Boyacá
Guata, Pmo. de
3650
Name
\0 \0
..::::;
See Pmo. de Gorgua
Guargua, Pmo. de
Cord. Oriental, ca. 40 km NE ofBogotá, ESE ofZipaquirá, S of Nemocón, E ofGuatavita and Guasca
Boyacá (?) Tibasosa
Cord. Oriental
Guerrero, Pmo. (de) (1)
3749
05°04'N
74°06'W
Cundinamarca
Tausa, Pacho, Zipaquirá & Cogua
Cord. Oriental, ca. 58 km N ofBogotá, NW of Zipaquirá, incl. Monte Grande; al so cal\ed Pmo. Guandoque
Guerrero, Pmo. de (2)
3400
07"47'N
72°58'W
N Santander
Salazar & Cáchira
Cord. Oriental, 1O km E of Cáchira; also called Ensillada de Guerrero
Güina, Pmo. de
3200
06°10'N
72°50'W
Boyacá
Belén
Cord. Oriental, between Belén and Susacón; also called Pmo. del Güina
Aracataca
Guática, Pmo.
Boyacá
Guircanapunanariga, Cerro
4500
Magdalena
Hatico, Pmo. de(!)
2900
N Santander
Hermosas, Pmo. (de) Las
4200
03°44'N
75°56'W
Valle/Tolima
Herveo, Pmo. de
4300
05°05'N
75°20'W
Caldas/Tolima
Hinchadero, Pmo. El
3700
Hirva, Pmo. de
3800
05°28'N
72°51'W
Hondón, Cerro
4000
Horqueta, La
5775
Hoya Hemando, Pmo.
3600
Sierra Nevada de Santa Marta Cord. Oriental, between Toledo and Pamplona
Buga & El Cerrito/ Chaparral
Cord. Central, N ofPmo. de La Estrella, N ofPmo. El Chinche and connected with it, NE ofPalmira, E ofBuga
..,
>
~ ;.;:: o
(/J
Cord. Central; also called Pmo. Aguacatal Cord. Central, Volcán Puracé area
Cau ca Boyacá
Recetor
Cord. Oriental, SE ofLaguna de Tota
Nariño
Cumbal
Nudo de Los Pastos
Magdalena
Aracataca & Santa Marta Sierra Nevada de Santa Marta, composed ofPico Cristóbal Colón and Pico Simón Bolívar
l0°50'N
73°41'W
Cundinamarca/Meta
Fómeque/El Calvario
Cord. Oriental, 1 km W ofLaguna Hoya Hernando
72°22'W
Boyacá
Aquitania (?)
Cord. Oriental; does this equal Hoya Verde?
Hoyevera, Pmo. de
4200
06°23'N
Huecas, Pmo. de
3400
06°10'N
72°25'W
Boyacá
Huila, Nevado del
5750
03°0l'N
76°00'W
Cauca/Huila/Tolima
Huila, Pmo. de
3500
04°56'N
73°45'W
Cundinamarca
Cord. Oriental, NE ofBogotá, NE ofGuasca, E ofGuatavita
Huila, Pmo. del
3700
02 o59'N
75°59'W
Huila
Cord. Central, on Nevado del Huila (peak at 5750 m), W side along Belalcázar-Cali rd.
Huina, Pmo. de
Cord. Oriental, SE ofChita, E side ofLaguna Chucas Páez/Teruel/Planadas
Cord. Central, E ofSantanderde Quilichao, SW ofPlanadas, W ofNeiva
See Pmo. de Güina
v; V,
N ame
Elev. (m)
Latitude
Longitud e Division 1
Division2
Notes Sierra Nevada de Santa Marta
Icayamagaca, Alto de
4000
Magdalena
Aracataca
Iguaque, Pmo. de
3800
05°40'N
73°27'W
Boyacá
Tunja, Arcabuco & Villa Cord. Oriental, witbin Santuario Flora y Fauna Iguaque, S of de Leyva Arcabuco and NE ofVilla de Leyva
Imbula Chico
3400
00°56'N
77°37'W
Nariño
Pupiales
Imbula Grande, Pmo.
U> 0\
Cord. Occidental, N oflpiales See Pmo. Rucio
India, Pmo. La
3800
04°07'N
75°43'W
Tolima
Cord. Central, along border witb Quindío depto., NNW of Roncesvalles
Iraca, Pmo. de
4200
03 o 13'N
76°04'W
Cauca!V alle/Tolima
Cord. Central, SSE ofPalmira, E ofPuerto Tejada; also called Pico Iraca
Jerusalén, Cerro
4000
Boyacá
Chita
Cord. Oriental
Jordán, Pmo.
3400
Santander
Tona
Cord. Oriental
~ (/)
Juan Rodríguez, Pmo. de
3400
72°58'W
Santander
Tona & Piedecuesta
Cord. Oriental, SW ofBerlín
Juanoy, Cerro de
3600
76°48'W
Nariño/Putumayo
Sibundoy
Cord. Centro-Oriental, N ofSibundoy; incl. Cerro Cascabel
o ,.., ""
07"08'N 01 o24'N
Junguilla, Pmo. de
See Pmo. de Yunguilla
Juntas, Pmo. de
3300
Juntas, Pmo. de Las
3600
00°48'N
77°26'W
Nariño
Juradó, Pmo. de
3160
06°54'N
72°39'W
Santander
Jurisdicciones, Pmo. de Las
3800
07"50'N
73°l5'W
N Santander/Cesar
·cord. Central, an extension ofPmo. de Guanacas at Km 53
Cauca Potosí
Cord. Oriental, S ofPmo. Almorzadero and connected witb it, N ofMálaga Abrego & Cáchira/Río de Oro
Antioquia
Laguna, Pmo. La Laguna, Pmo. de La
3750
Lajas, Pmo. Lajita, Pmo.. La
3400
05°59'W
72°34'N
3100 74°48'W
Cord. Oriental, SW ofOcaña; coordinates approximate Cord. Central, NW ofMedellín, part ofthe Belmira system
o
::r::
tr1
~
::.8 ~
o ~ ti:J o ,..,
z~
o ~
r
Caldas
Cord. Central, W slopes, on Nevado del Ruiz
o
Boyacá
Cord. Oriental; also called El Perro
ES
Boyacá 02°54'N
SW of Pmo: Guapuscal and connected witb it, E oflpiales, S of Pmo. El Palacio and connected witb it; also called Pmo. Las Juntas
2:: tr1 2::
Monguí
Cord. Oriental Cord. Oriental, E ofNeiva; also called Cord. de Los Picachos
Leiva, Cerro
3520
Meta
Granada
León, Cerro ( 1)
4000
Boyacá
Chita
Cord. Oriental
León, Cerro (2)
3900
La Guajira
Riohacha
Sierra Nevada de Santa Marta
Leona, Pmo. La
3800
04°04'N
74°04'W
Cundinamarca/Meta
Cord. Oriental, W ofVillavicencio
Leona, Pmo. Monte de
3200
05°27'N
73 o lO'W
Boyacá
Cord. Oriental, SE ofTunja, SW ofLaguna de Tota; also called Pmo. del Carmen
Leonera, Pmo. La
3800
03°46'N
75°49'W
Tolima
Cord. Central, SSE ofSanta Lucía, NNW ofRJo Blanco
Letras, Pmo. Las
3700
Caldas
Cord. Central
~
ztr1
<o r
00
.¡:.
N ame
Elev. (m)
Letrero, Pmo. (de)!
3688
Llano Grande, Pmo. de
Latitud e 01 °53'N
Longitude 76°32'W
Division 1
Division 2
Notes
Huila/Cauca
San Agustín/San Sebastián
Macizo Colombiano, NW ofPmo. Cutanga and connected with it, S ofPmo. de Las Papas and connected with it; S ofPopayán
3750
Boyacá
Monguí
Cord. Oriental
. Llanos de Cuibá
2850
Antioquia
Santa Rosas de Osos
Cord. Central; also spelled Cuivá; subpáramo
Loma El Páramo
3200
01 °06'N
7r33'W
Nariño
Loma Grande, Pmo. de
3600
or29'N
72°57'W
N Santander
Loma Larga
3400
00°57'N
7r37'W
Nariño
Loma Paramillo
2600
01 °07'N
7r30'w
Nariño
Loma Puerta del Páramo
2900
01 °l8'N
7rl7'W
Nariño
Loriana, Pmo. de La
2750
00°57'N
7ro4'W
Nariño
Pasto
S ofLago Guamués
Lúcura, Pmo. de
3200
orJJ'N
72°57'W
Santander
Tona
Cord. Oriental, W ofBerlín
Cundinamarca
Cord. Oriental, Usme-San Juan rd., near side rd. to Nazareth
Magdalena
Sierra Nevada de Santa Marta, near Pmo. de Chirugua
WNW ofTúquerres; probably not páramo Cord. Oriental, WSW ofCucutilla Pupiales
Cord. Occidental, N oflpiales WNW ofTúquerres, E of Guaitarilla; probably not páramo N ofPasto, along W si de of rd. to airport; probably not páramo
Lumapán, Pmo. de
3200
Macatama, Pmo. de
4570
10°5l'W
73°4l'N
Cundinamarca (?)
See map in Hettner, 1892; and Guhll966: 62, 100
Macotama, Pmo. de
3800
10°56'N
73 °37'W
La Guajira
Sierra Nevada de Santa Marta; probably the same as Pmo. de Macatama, and also spelled Macotoma
Mamapacha,Pmo. de
3200
Mamarongo, Pmo. (de)
3500
10°55'N
73 °20'W
La Guajira
Sierra Nevada de Santa Marta, NE slopes, E ofMacotama
Mamito, Pmos. de Laguna
4300
Magdalena
Sierra Nevada de Santa Marta, below and SW of Picos Reina and Ojeda
Mamo, Pmos. de Laguna
4300
Magdalena
Sierra Nevada de Santa Marta, below and SW ofPicos Reina andOjeda
Machetá, Pmo. de
Boyacá
Garagoa Chinavita
Cord. Oriental, SE ofTunja
06°54'N
72°45'W
Santander
Cord. Oriental; also called Pmo. Blanco (2)
Marchán, Pmo. de
3424
05°40'N
73°42'W
Boyacá/Santander
Cord. Oriental, N ofLaguna de Fúquene, E ofChiquinquirá
Marruecos, Pmo.
3600
04°08'N
75°37'W
Tolima
Cord. Central, N ofRoncesvalles
Martino, Pmo.
3300
Santander
Cord. Oriental
Mata, Alto de La
4000
06°l2'N
72°5l'W
Santander/Boyacá
Cord. Oriental, NNE ofDuitama, WSW ofCharalá
Matarredonda, Pmo.
3000
05°20'N
74°03'W
Cundinamarca
Cord. Oriental, E ofLa Palma, SW ofLaguna de Fúneque
Méndez, Pmo. de
4400
06°36'N
72°22'W
Boyacá
Cord. Oriental, NE ofLas Mercedes; also called Pmo. de Contento
Mercedes, Pmo. Las
3600
04°06'N
74°00'W
Cundinamarca/Meta
Cord. Oriental, W ofVillavicencio
Mantilla, Pmo.
Mesa Alta, Pmo. de
'D 'D
~
"" > ~ ;s:: o
C/)
See Pmo. de Mesalta
v; -..)
Name
Elev. (m)
Latitud e
Longitud e Division 1
Division 2
Notes
Mesalta, Pmo. de
3800
05°56'N
72°44'W
Tasco
Cord. Oriental, NE ofDuitama, NE ofTasco, S ofPaz de Río; also caiied Pmo. de Mesa Alta
Mina, Cerro
3750
08°l8'N
73o li'W
N Santander
Cord. Oriental, SE ofLa Convención, NE ofOcaña
Miraflores, Pmo. de (1)
3600
03 °55'N
75°56'W
Vaiie/Tolima
Cord. Central, N ofPmo. de Las Hermosas and connected with it, S of Pmo. Barragán and connected with it, ESE of Tuluá, NW of Chaparral
Miraflores, Pmo. de (2)
3300
Mirlas, Pmo. de Las Mogotocoro, Pmo. de
3800
Boyacá
Huila/Caquetá
Cord. Oriental, near Gigante
Chocó
Not true páramo
Santander
Cord. Oriental, near Las Vetas; sometimes speiied Mogorotoco
V> 00
s;: tTl s;: o
Monsalve, Pmo. de
3600
or25'N
72°54'W
Santander
Monserrate, Pmo. (de)
3250
04 °37'N
74°04'W
Cundinamarca
Monte de Leona, Pmo.
3200
05°27'N
73°l0'W
Boyacá
Monte de Luz, Pmo.
3000
Monterredondo, Pmo.
3500
Moras, Pmo. de (Las)
3800
Morro Frontino, Pmo. del
3900
Morro Paramillo
3000
01 °l5'N
77°34'W
Nariño
Morro, Pmo. El
3130
06°35'N
75°32'W
Antioquia
Morrobravo, Pmo. de
3400
07"05'N
72°55'W
N Santander
Cord. Oriental, N of Santa Bárbara, SW of Berlín
> z ñ > l'
Mortiño, Pmo. (del)
3700
N Santander
Cord. Oriental, between Cúcuta and Bucaramanga
Cl
Moscopán, Pmo. de
3400
Cauca
Cord. Central, 32 km from Puracé on rd. to Moscopán
Mosquito, Pmo. de Laguna del
3850
Caldas
§ zlTl
02°50'N
76°l2'W
Cord. Oriental, NNE ofCalifornia Bogotá
Cord. Oriental, ca. 7 km NE of Bogotá; also caiied Cerro Monserrate Cord. Oriental, SE ofTunja, SW ofLaguna de Tota; also cailed Pmo. del Carmen
Boyacá
Saboyá
Cord. Oriental
Cauca
Silvia & Paéz
Cord. Central
Cauca
Coid. Central, NNE ofPopayán, between Mozoco and Pitayó,
Antioquia
Cord. Occidental, N ofUrrao
SW ofPmo. de Santo Domingo and connected with it W ofPasto, NE ofTúquerres; not páramo Belmira
Cord. Central, part ofPmo. de Belmira, N ofMedeiiín
Cord. Central, source ofRío Otún, below Nevado de Santa Isabel
Mucute, Pmo. de
...:¡
::e:
tTl
ztTl
::E o"<
~ IJ:j o...:¡
>
Nariño
Mundo Nuevo, Pmo.
See Pmo. Nuevo Mundo
Mundonuevo, Pmo.
See Pmo. Nuevo Mundo 3850
Muzo, Pmo. de N aboba, Pmos. de Laguna
o'Tl
From type of Acrostichum calaguala (Moritz 315)
Muerte, Pmo. de la
Musinga, Alto
;;:;<Zl
06°39'N
76°40'W
Antioquia Boyacá
4300
Magdalena
Cord. Occidental, S ofFrontino, N ofUrrao; páramo ? Muzo
Cord. Oriental, W of Chiquinquirá Sierra Nevada de Santa Marta, below and SW ofPicos Reina and Ojeda
<o r
00 .jo.
N ame
Elev. (m)
Nabulgalha, Pmo. de
3350
Nariz, Pmo. de La (1)
3300
Nariz, Pmo. de La (2)
Latitud e
Longitude Division 1
Division 2
Magdalena Boyacá 10°40'N
73°20'W
Cesar
Notes Sierra Nevada de Santa Marta
Santa Rosa Sierra Nevada de Santa Marta, N ofValledupar
Negro, Cerro (1)
4470
00°5l'N
7r55'W
Nariño
Cumbal
Cord. Occidental, W oflpiales
Negro, Cerro (2)
4276
01 °44'N
7r36'W
Nariño
Ipiales & Potosí
Cord. Centro-Oriental, E oflpiales
Cundinamarca (?)
Neusa, Pmo. Nevaditos, Pico
5375
Nevado, Pmo. El
4000
03°56'N
74°!0'W
Meta
Cord. Oriental, part ofPmo. de Sumapaz (?), W ofCubarral, S ofPmo. El Gallo; also called Cerro El Nevado or Alto Nevado
Nieves, Pmo. Las
3900
03°48'N
75°49'W
Tolima
Cord. Central, SSE ofSanta Lucía, NNW ofRío Blanco, N of Pmo. La Leonera
Tolima
Roncesvalles
Cord. Central
04°!8'N
74°05'W
Cundinamarca
Une & Gutiérrez
Cord. Oriental, NE of Pmo. Sumapaz, SW of Cáqueza; al so called Pmo. Mundonuevo or Pmo. Mundo Nuevo
Norrnandía, Pmo. de
3200
Nuevo Mundo, Pmo.
3500
Sierra Nevada de Santa Marta
Obelisco, Pmo. del
3200
Cundinamarca
Ocetá, Pmo.
3800
Boyacá
Ocungahla, Pmo. de
3690
Cord. Oriental, near Pacho Monguí
See Pmo. Ocusá Boyacá
3000
Paipa
'1:!
>~ ~ oCll
Cord. Oriental; also called Pmo. Ocusa de Blanco
Boyacá
Ogonta, Pmo. de Ojeda, Pico
Cord. Oriental; also spelled Osetá Sierra Nevada de Santa Marta
Ocusa de Blanco, Pmo. Ocusá, Pmo.
'-0 '-0
.::S
5490
Ojo, Pmo. de Olleta, Cráter (de) La
4750
10°50'N
73°38'W
Magdalena/Cesar
06°55'N
72°45'W
Santander
04°5l'N
75°23'W
Caldas/Tolima
Sierra Nevada de Santa Marta, NW ofValledupar Cord. Central, NW ofNevado del Ruiz, SE ofManizales; also called Cerro O lleta
Orocué, Cerro de
3900
Oseras, Alto (de las)
3830
03°45'N
74°30'W
Cesar/N Santander
Cord. Oriental, S of Abrego; also spelled Oroque
Huila!Meta/Cundinamarca
Cord. Oriental, 50 km SE ofEspinal, S end ofPmo. de Sumapaz
Osos, Pmo. de Los
4000
06°!9'N
72°!9'W
Arauca
Otún, Laguna del
3800
04°47'N
75°26'W
Risaralda
Pereira
Ovejas, Pmo. de Las
3500
01 o IO'N
7rl4'W
Nariño
Pasto
Serranía to the ESE of Lago Guamués
Ovejeras, Pmo. (de las)
3000
Ü5°]]'N
73°46'W
Cundinamarca
Suesca
Cord. Oriental, NNE ofSuesca, E ofLaguna de Suesca
Páez, Pmo. de La Laguna de(!)
3450
Cauca
Cord. Oriental, Sierra Nevada del Cocuy Cord. Central, 2 km W of summit ofNevado de Santa Isabel
Cord. Central, W slopes, headwaters ofRío Páez
:::;; '-0
Elev. (m)
Latitude
Longitude
Division 1
Division2
Notes
Paja Blanca, Pmo. de
3400
00°02'N
7r33'W
Nariño
Guachucal, Ipiales, Ospina & Pupiales
Nudo de Los Pastos, actually extends from 00 °02'N toO 1 o o 1'N and 77°33'W to 7r39'W; also spelled Pajablanca
Palacio, Pmo. (de El)
3700 00°38'N
1r2ow
Nariño
Ipiales & Potosí
Cord. Centro-Oriental, between La Victoria and Monogamba, SE of and running parallel to Pmo. de Las Juntas, continues with Cerro Palacio and Cerro San Francisco; also called Pmo. Palacios
N ame
3500
Paletará, Pico
4482
02o!O'N
76°26'W
Cauca!Huila
Palomas, Pmo. de Las
3290
05°45'N
75°20'W
Antioquia
Pamplona, Pmo. de
3500
Pan de Azúcar, Cerro
4670
Pan de Azúcar, Pmo. (1) Pan de Azúcar, Pmo. (2)
Cord. Central, SE ofPopayán, WSW ofLa Argentina Sonsón
3500
05°57'N
3600
03°48'N
76°19'W
73°02'W 76°05'W
Cauca/Huila
Boyacá
Cord. Central, Vereda San Francisco, near Sonsón Cord. Oriental, W ofMutiscua; refers to Pmo. de Santurbán
N de Santanter 02°13'N
Puracé!La Argentina
Cerinza!Duitama
Cord. Central, Sierra Nevada de Los Coconucos, SE ofPopayán, W ofLa Argentina; also called Volcán Pan de Azúcar and Cerro Agua Blanca Cord. Oriental, S ofPmo. de la Rusia and connected with it Cord. Central, NNE ofPalmirá, N ofPmo. de Los Domínguez
Valle
See Cerro Pan de Azúcar
Pan de Azúcar, Volcán 4200
01 °55'N
76°36'W
Cauca
Paramillo (1)
3000
02°55'N
76°24'W
Cauca
Cord. CentraC ESE ofSantander de Quilichao; perhaps not true páramo
Cundinamarca
Cord. Oriental; a town not páramo
2200
06°00'N
73°05'W
N Santander
Cord. Oriental, WSW of Abrego; name for school and few houses, not páramo
Paramillo (3)
San Sebastián
Macizo Colombiano, S ofPopayán, 70 km W ofPitalito, 40 km W ofSari-Agustín
Papas, Pmo. (de) Las
Paramillo (2)
See Nudo de Paramillo
Paramillo del Sinú
o
Cord. Oriental, 30-40 km ENE ofBogotá, region ofChingaza
Cundinamarca
Palacio, Pmo. El
0\
Paramillo, Alto El
1500
08°37'N
73 °25'W
N Santander
Cord. Oriental; not páramo
Paramillo, Cerro
3000
02°32'N
75°12'W
Huila/Caqueta
Cord. Oriental, SSE ofNeiva, E of Algeciras
Paramillo, Llano de
1800
06°0l'N
73°05'W
N Santander
Cord. Oriental, WSW of Abrego; not páramo
Paramillo, Loma (1)
2600
01 °07'N
7r3ow
Nariño
E of Guaitarilla and WNW ofTúquerres; probably not páramo
Paramillo, Loma (2)
3600
04°00'N
75°55'W
Valle
Cord. Central, SW ofBarragán, NE of Santa Lucía
Paramillo, Nudo de
3960
oro4'N
75°55'W
Antioquia
Cord. Central, NE ofDadeiba, W ofHuango; also called Cerro Paramillo or simply Paramillo
Paramito, Hacienda
1650
06°35'N
73°1l'W
Santander
Cord. Oriental; a hacienda NNW of San Gil; not páramo
Páramo (1)
1300
06°31 'N
73 °2Ü'W
N Santander
Cord. Oriental, N of Simacota; local name for a group of homes, not páramo
~ rrJ ~
o
~
m
o ..., "rj
;:¡::
rrJ
zrrJ
:E >-<: o
~ to o..., >~
(J
>1:"' o >-
ElrrJ z
< ~
00 ~
N ame
Elev. (m)
Latitud e
Longitude
Division 1
Division2
Notes
Páramo (2)
1000
Nariño
A town, not páramo; old name for town ofRicaurte
Páramo (3)
3475
Santander
Cord. Oriental, Motocoro, 2 km S ofVetas
Páramo (4)
1400
06°25'N
73°10'W
Santander
Cord. Oriental, town S ofSan Gil; not páramo
Páramo (El)
3000
05°!8'N
75°2l'W
Caldas
Cord. Central, SE ofPalamina, NE ofManizales
Páramo, Alto El
1600
05°38'N
72°25'W
Boyacá
Cord. Oriental, 20 minutes (oflatitude) N ofYopa!; not páramo
Páramo, Cerro (1)
2000
08°24'N
73°14'W
N Santander
Cord. Oriental, S of San Calixto, NE of Ocaña; not páramo
Páramo, Cerro (2)
1600
06°25'N
73°23'W
Santander
Cord. Oriental, WSW ofSimacota; not páramo
Páramo, Cerros del
1200
05°38'N
7Z025'W
Boyacá
Cord. Oriental, 20 minutes (oflatitude) N ofY opa!; not páramo
Páramo, Cuchilla El
1900
05°40'N
72°25'W
Boyacá
Cord. Oriental, mountain range N of Yopa!
Páramo, El (1)
3000
05°35'N
73°55'W
Boyacá
Cord. Oriental; town WSW of Chiquinquirá; not páramo
Páramo, El (2)
1700
08°24'N
73°l4'W
N Santander
Cord. Oriental; name ofschool and group ofhomes; not páramo
Páramo, Loma El (1)
2000
or53'N
73°16'W
N Santander
Cord. Oriental, S of Abrego; not páramo
Páramo, Loma El (2)
3200
01 °06'N
7r33'W
Nariño
WNW ofTúquerres; probably not true páramo
Parroquia Vieja, Pmo. de
2900
Boyacá
Cord. Oriental
Pasea, Pmo. de
3800
Cundinamarca
From type specimen of Baccharis pascensii (Stuebel 133, F neg. 15025)
Pasquilla, Pmo.
3150
Cundinamarca
Pastora, Pmo. de La
3150
Nariño
Pastos, Nudo de los Patascoy de Santa Lucía, Cerro
4000
01 °00'N
7r3o'W
Nariño
00°57'N
7r04'W
N ariño/Putumayo
"<:!
> ~ S:: o tzl
15 km E ofPasto a!ongrd. to Mocoa; a!so cal!ed Pmo. de Tábano Incl. Volcán Chiles, Nevado del Cumbal, Cerro Alcalde, and Cerro Patascoy de Santa Lucía; also called Macizo de Pasto
Pasto/Santiago
SSE of Laguna de La Cocha, SE ofPasto; also called Cerro Patascoy and Volcán Patascoy See Cerro El Bordoncillo
Pataseos, Volcán Arauca
Cord. Oriental, Sierra Nevada del Cocuy, E slopes, an area of páramos ranging 4200-4300 m
1r26'W
Nariño
SE Nariño, on Ecuador border
72°50'W
Boyacá
Cord. Oriental, S ofPmo. Puchicavo and connected with it, E of Laguna de Tota, SE of Sogamoso
Patio Bolos, Pmos. de
4300
Pax, Cerro
3350
00°23'N
Pedrisco, Pmo. del
3500
05°35'N
Pelao, Cerro (El)
4000
Boyacá
Pelao, Cerro El
4000
Boyacá Source ofRío Caqueta
Peña Alta, Pmo. de Peña Lisa, Pmo.
Cord. Oriental, vicinity of Cabras Pasto
'D 'D
.:S
3500
04°06'N
73°58'W
Cundinamarca!Meta
Cord. Oriental, W ofVillavicencio, E ofPmo. Las Mercedes; also spel!ed Peñalisa
0\
N ame
Elev. (m)
Latitude
Longitude Division 1
Division 2
Notes
o,
N
Peña Negra, Pmo. de (1)
3600
Boyacá
Cord. Oriental, continuation ofPmo. de Gachaneque
Peña Negra, Pmo. de (2)
3625
04°47'N
73°49'W
Cundinamarca
Cord. Oriental, N ofCordillera Verde, NE ofBogotá, W side of high plain ofBogotá, near Subachoque
3800
03 °56'N
75°50'W
Valle
Peñas Blancas, Pmo. de
3600
04°42'N
75°25'W
Tolima
Peñitas, Pmo. (Las)
3700
Boyacá
Peñón, Pmo. del
3100
Cauca
Near Jambaló
Perijá, Sierra de
3350
Magdalena
Incl. Cerro Venado and Cerro Avión
o
See Pmo. Lajas
CZl
See Pmo. de Peña Negra(!)
Peñanegra,Pmo.de Peñas Blancas, Pmo.
Cord. Central, E of Santa Lucía, NW of Pmos, La Soledad, Diamante, and Rancho Largo Cord. Central, NW ofNevado del Tolima Monguí
Perro (El)
Cord. Oriental
~
m
~
~
Pesca, Pmo. (La)
3400
Boyacá
Tuta
Cord. Oriental
o'T1
Pescadero, Pmo. de
3400
07"12'N
72°54'W
Santander
Tona
Cord. Oriental, NW ofBerlín
::r::
Petacas, Pmo. de
4050
01 °34'N
76°46'W
Nariño/Cauca
La Cruz/Santa Rosa
NE ofPasto, ENE ofLa Cruz, on Volcán Petacas, continuation of Cerro A?imas; al so called Cerro Petacas
zm
Picacho, Cerro
4700
Pilar, Pmo. del Piñuela, Pmo. La
3000
Piranchón, Pmo.
3700
04°l2'N
74°16'W
Cundinamarca
02°05'N
74°10'W
Cundinamarca Boyacá
Pirineos, Pmo.
3600
04°l2'N
75°36'W
Tolima
Pisba, Pmo. de
3600
05°53'N
72°37'W
Boyacá
Piscicultura, Pmo. La
3400
01 °l0'N
77°05'W
Nariño
-3
m ~ >-<
o
Cord. Oriental, N of La Pradera, S of Pacho, NW edge of Sabana de Bogotá Zetaquira & Tota
Socotá
Cord. Oriental
z>-
Cord. Central, N ofRoncesvalles, N ofPmo. Marruecos, E of Geno va
ñ > r
Cord. Oriental, NE ofSogamoso, N ofPmo. de San Ignacio and connected with it; also spelled Pisva
>-
E of Pasto, NE of Laguna La Cocha; also called Pmo. del Quilinsayaco
Pisva, Pmo. de
~
to
o-3
o
1:1 ""'m z
See Pmo. de Pisba
Plan del Cóndor, Pmos. de
3970
Planos del Toldadero
3950
Caldas
Cord. Central, below del Nevado de Santa Isabel, headwaters of Río Otún
Boyacá
Plateado, Cerro
3250
02°20'N
Playón, Alto El
3500
06°l9'N
77°l5'W
Cauca
72°!9'W
Arau ca
Balboa/Argelia
Cord. Occidental, along Balboa-Argelia rd. at the crest ofthe cordillera and headwaters o fRío Plateado and Río Guaitara Cord. Oriental, Sierra Nevada del Cocuy
<o !:"
00
.p.
N ame
Elev. (m)
Portachuelo, Cerro
4400
Potosí, Cerro
3800
Potrogaca, Pico
4350
Latitude
Longitud e
Division 1
Division 2
'D 'D
Notes
~
01 °27'N
76°50'W
00°53'N
7r24'W
Nariño
Córdoba & Puerres
Cord. Centro-Oriental, ENE oflpiales, near Las Juntas
Sal azar
Cord. Oriental, S of Pmo de Guerrero an connected with it, WSW of Cúcuta, SW of Salazar
Nariño/Putumayo
El Tablón!Sibundoy
N Santander
ENE ofPasto Cord. Oriental, around Ocaña; not true páramo
Pozo Alto, Pmo. de
1800
Precipicio, Pmo. del Cerro
3600
Pregonero, Alto (El)
4000
Pringador, Pmo. de
3400
or44'N
72°57'W
N Santander
Puchicavo, Pmo.
3500
05°40'N
72°45'W
Boyacá
Cord. Oriental, E of Somagoso, S of Pmo San Ignacio and connected with it; also called Pmo. Puchicaro
Puentes, Pmo. (de) Los
3900
07°22'N
72°52'W
Santander
Cord. Oriental, abo ve La Baja, NE of California, N of Vetas; probably same as Pmo. de Las Puentes
Puerta del Páramo, Loma
2900
Ol 0 !8'N
7rl7'W
Nariño
N ofPasto, along rd. to airport; probably not páramo
Puertas, Pmo. de La
3700
Púlpito del Diablo, Cerro
5187
06°22'N
Puchicaro, Pmo.
See Pmo. Puchicavo
72°20'W
Santander
Cord. Oriental, close to La Baja
Boyacá/Arau ca
Cord. Oriental, Sierra Nevada del Cocuy
Puracé, Pmo. de(!)
4400
02°24'N
76°23'W
Cauca!Huila
Quilinsayaco, Pmo. de
3200
01 °08'N
7ro7'w
Nariño/Putumayo
Quindío, Pmo.
4300
04°38'N
75°25'W
Caldas/Tolima
Rabanal, Pmo.
3590
05°25'N
73°35'W
Boyacá
Rabón, Pmo. del
3000
05°l4'N
74°08'W
Cundinamarca
Rabona, Alto (de)
4082
04°05'N
74°13'W
Cundinamarca
Ramal, Pmo.
2800
'"O
> > ;::::
;;¡;¡
Cord. Central, slopes ofVolcán Puracé (4800 m), ESE ofPopayán Pasto/Santiago
o
Pasto-Mocoa rd., Km 35 E of Pasto, between El Encano and Santiago; also known as Pmo. de La Piscicultura; also spelled Quilinchayaco (?)
C/)
Cord. Central, slopes ofNevado (del) Quindío (5150 m), ESE ofPereira, ENE ofSalento, NNW oflbagué
Quitasol, Pmo.
See Cerro Altar
Cundinamarca
V entaquemada Samacá
Cord. Oriental, ESE ofLaguna de Fúquene, SW ofTunja, N of Pmo. Albarracín and connected with it Cord. Oriental, SE ofLa Palma, N ofPacho Cord. Oriental, W border ofPmo. Sumapaz, E ofEspinal
Suesca
Cord. Oriental; probably not páramo
Ramírez, Alto (de)
3788
03°49'N
74°2l'W
Cundinamarca!Meta
Cord. Oriental, part ofPmo. de Sumapaz
Ramírez, Pmo.
3800
or29'N
7ZC54'W
Santander
Cord. Oriental, E of Carchiri
Ramos, Cerro
4200
Rancho Largo, Pmo.
3800
03°55'N
75°50'W
Valle
Cord. Central, ESE ofSanta Lucía, W ofPmo. La Soledad and Pmo. Diamante
-"' w
N ame
Elev. (m)
Latitud e
Longitude
Division 1
Division 2
Rayo, Cerro El
3720
03°5l'N
74°20'W
Cundinamarca!Meta
Cord. Oriental, on Pmo. de Sumapaz
Rechíniga, Pmo. de
5223
06°l6'N
72°23'W
Arauca/Boyacá
Cord. Oriental, S part ofSierra Nevada del Cocuy, E ofSusaeón; also called Pico de Rechíniga and Farallones de Rechíniga
Reina, Pico La
5535
Magdalena
Sierra Nevada de Santa Marta
Remedios, Alto de Los
Notes
0\
_¡,.
Macizo Colombiano, high point in Pmo. de Las Papas
Rico, Pmo.
4200
07° 18'N
72°55'W
Santander
Cord. Oriental, NE of Bucaramanga, E of Charta, SSW of Vetas, NW ofPmo. Santurbán and connected with it
Rincón, Pmo. El
4000
06°24'N
72°22'W
Boyacá
Cord. Oriental
Río Sevilla, Pmo. de
3410
Magdalena
Sierra Nevada de Santa Marta
Riosucio, Pmo. de
3225
Cauca
Ritacuva, Alto (de)
5493
Inza
Cord. Central, along Totoró--Inza rd.
;;;:: tT1 ;;;::
o
;:a
06°30'N
72°19'W
Boyacá
Cord. Oriental, Sierra Nevada del Cocuy; aiso spelled Ritacuba
03 °47'N
75°57'W
Valle
Cord. Central, ENE of Albania, N ofPmo. de Las Hermosas
.,o
Cord. Oriental; not páramo
::r:
(/J
Rocío, Pmo. (d)el
4000
Rodríguez, Pmo.
2400
Romeral, Pmo. de(!)
3800
06°40'N
72°l8'W
Boyacá
Romeral, Pmo. de (2)
3700
or23'N
72°52'W
Santander
Rosal, Pmo. del
3700
00°52'N
1r22'W
Nariño
Puerres
Rucio, Pmo.
3400
01 °58'N
7r34'w
Nariño
El Contadero, Gualrnatán, Córd. OcCidt;ntal; also called Pmo. Imbula Grande Ospina & Sapuyes
(';
Rucio, Pmo. (del)
2800
Huila/Meta
Colombia/Granada (Boca de Monte)
>-3
Ruiz, Pmo. de
3500
Bolívar
04 °54'N
75°l8'W
Jenesno
· Cord. Oriental, Sierra Nevada de Güicán
Caldas/Tolima
Ruiz, Nevado del Runta, Pmo. de
3278
05°30'N
73 °25'W
Boyacá
Rusia, Pmo. (de La)
4320
Ruso, Pmo.
3150
Santander
Sábana Rubia, Pmo. de (2)
3250
Magdalena
Sabanazo, Pmo. de
3280
05°59'N
73°05'W
Sábana Rubia, Pmo. de (1)
Santander/Boyacá
Paipa & Duitama/Santa Rosa de Viterbo
>-3 tT1
ztT1
Cord. Oriental, Hoya del Río Cucutilla, NE ofCalifornia, N ofV etas
::E
Cord. Centro-Oriental, E ofPuerres, rd. to Monopamba
o--<
Cord. Oriental; probably not páramo
"'o
to
z>-
Cord. Central, ón slopes ofNevado del Ruiz (5432 m), N of !bagué, SE ofManizales
ñ >r'
See Pmo. de Ruiz
>-
Cord. Oriental, SW of Tunja, NE of Pmo. Gachaneque and connected with it
E3
Cl
tT1
z
Cord. Oriental, N ofPmo. Pan de Azúcar and connected with it, 21 km NNW ofDuitama; may incL Pmo. de Agüero
Cesar/Magdalena
06°35'N
75°32'W
Antioquia
Sierra de Perijá, Colombia/Yenezuela border area, part of Cerro del Avión (Cerro Monte Plano in Venezuela) Belmira
Cord. Central, part of Altiplano de Santa Rosa de Osos, part of Pmo. de Belmira; also called Pmo. de Alto de Sabanazos
<o r-'
00
_¡,.
N ame
Elev. (m)
Latitude
Longitud e Division 1
Division 2
Sabanazos, Pmo. de Alto de
Notes See Pmo. de Sabanazo
05°45'N
Saboyá, Pmo. de
4003
73°5l'W
Salado, Pmo. (El)
3500
Salazar, Pmo.
3000
Salitre, Cerro El
3800
Salitres, Pmo. Las
3600
Cundinamarca
Cord. Oriental, S ofBogotá, ESE ofEmbalse Chisacá
San Antonio del Bordoncillo, Pmo. de
3250
Putumayo
Between El Encano and Sibundoy; also called Pmo. de San Antonio
San Antonio, Pmo. (2)
3350
Cundinamarca
San Cayetano, Pmo. de
3750
oso 18'N
74°04'W
Cundinamarca
San Félix, Pmo.(!)
3650
05°25'N
75° 19'W
San Félix, Pmo. (2)
3100
San Fernando, Cerro
3810
San Fortunato, Pmo. de
3000
San Francisco, Cerro
3685
or48'N
72°52'W
04°l8'N
74°07'W
Boyacá/Santander
Cord. Oriental, N of Chiquinquirá; al so called Peña de Saboyá
Valle
Buga
Cord. Central
N Santander
Sal azar
Cord. Oriental, WSW ofCúcuta, NW ofSalazar, E ofPmo. de Cáchira and connected with it
Boyacá
San Antonio, Pmo. (1)
See Pmo. de San Antonio del Bordoncillo
San Cristóbal, Pmo. de
Cord. Oriental, 2 km NNW ofLaguna de Chingaza San Cayetano
Cord. Central; coordinates are for San Cayetano
Caldas
Pensilvania
Cord. Central, E ofRiosucio and Salamina, W of Cordillera de Miraflores, S ofPmo. Arboleda and connected with it
Antioquia
Bello
Cord. Central, 30-35 km NW ofMedellín along rd. toward San Pedro
Cundinamarca
05°35'N
00°48'N
76°03'W
7r29'W
San Francisco, Pmo. ( 1)
Antioquia/Chocó
Cord. Occidental, Farallones de Citará, SW of Andes, W of Jardín; also called Alto de San Fernando
Cundinamarca
Cord. Oriental, near Bogotá, extreme S rim ofBogotá tabeland; old name used by Goudot and Hartweg in tbe mid-1800s
Nariño
Córdoba & Potosí
3200
Putumayo
San Francisco
San Francisco, Pmo. de (3)
3900
Cauca
San Sebastián
San Ignacio, Pmo. de
3400
San José, Pmo. (1)
3800
Cundinamarca
San José, Pmo. (2)
3600
Meta
05°47'N
72°42'W
San Miguel, Pmo.
San Pablín, Laguna de
"'> ~ 2:: o
(/)
E oflpiales, continuation ofCerro Palacio
Cundinamarca
San Francisco, Pmo. (2)
San Nicolás, Cerro (de)
\0 \0
::S
3780
05°40'N
76°05'W
Boyacá
Cord. Oriental, NE of Somagoso, NE of Mongua, S and connected with Pmo. de Pisba Cord. Oriental, 3.5 km SE ofLaguna de Chingaza Calvario
Cord. Oriental
Cundinamarca
Cord. Oriental, above Hacienda Los Arenillos
Antioquia/Chocó
Cord. Occidental, Farallones de Citará, W of Andes, SW of Betania; probably not páramo
Boy;tcá
Cord. Oriental, Sierra Nevada del Cocuy o-,
'-"
N ame
Elev. (m)
Latimde
Longitude
Division 1
Division2
Cau ca
Totoró
07"47'N
73°03'W
N Santander
Cord. Oriental, partofPmo. de Cáchira, E ofCáchira, between Las Ventanas and Pmo. de Guerrero
Cauca
Cord. Central, small part of Pmo. de Puracé on the E slopes, headwaters ofRío San José
San Pedro, Pmo. de (1)
3225
San Pedro, Pmo. de (2)
3000
San Rafael, Pmo. de
3350
San Salvador, Pmo.
3600
04°44'N
73°47'W
Cundinamarca
Santa Inés, Pmo.
3000
06°48'N
75°43'W
Antioquia
Santa Lucía, Pmo. Santa Lucía, Pmo. de
Cord. Central, along Totoró-Inza rd.
Cord. Central, 80 km NW ofMedellín, NNE of Antioquia, E of Rodadas and Fortuna, part ofthe Belmira system; al so spelled Santa Inéz See Pmo. Santa Inés
Santa Isabel, Pmo. (de) (!) 3800
3680
07"20'N
72°20'W
04°47'N
75°26'W
03 °58'N
75°55'W
3100
Santa Maria, Nevado
N Santander
Toledo
Risaralda!Tolima
Valle
Cord. Oriental, within Parque Nac. Nat. Tamá, E ofToledo; coordinates approximate Cord. Central, slopes Nevado de Santa Isabel (5100 m), SE of Nevado del Ruiz, N ofNevado del Quindío, E ofPereira, NNW oflbagué, incl. Laguna de Otún
Tuluá
Putumayo
Cord. Central, SW ofBarragán, NE ofSanta Lucía, S ofLoma Paramillo, N ofPmo. Barragán; may equal Paramillo Santa Lucía S side Laguna de La Cocha, source ofRío Alisales
04°48'N
75°22'W
Tolima
!0°50'N
73°40'W
Magdalena/Cesar/La Guajira
Sierra Nevada de Santa Marta, NW of Valledupar, incl. Pico Cristóbal Colón, Pico Codazzi, Pico Simón Bolívar, Pico Simons, Pico Ojeda, and Pico El Guardián; equals Macizo de Santa Marta
Boyacá
Cord. Oriental, between Santa Rosa de Viterbo and Cerinza
Risaralda
Cord. Central, W slopes Nevado del Ruiz, E ofSanta Rosa de Cabal, ENE ofPereira
Santa Marta, Sierra Nevada de
5800
Santa Rosa, Pmo. de (1)
3000
Santa Rosa, Pmo. de (2)
4600
Santa Rosita, Pmo. (de)
3400
Santo Domingo, Pmo. (de)
4000
03°00'N
Santo Ecce Horno, Pmo. de
3500
Santurbán, Pmo. de
4030
Sarna, Pmo. de La Sátiva, Pmo.
04°50'N
75°30'W
a, a,
Cord. Oriental, E ofBogotá, N ofPmo. Las Barajas, SE of Guasca Belmira & San Andrés
Santa Inéz, Pmo.
Santa Isabel, Pmo. de (2)
Notes
Boyacá
Susacón
76°05'W
Cauca!Tolima
Toribío, Crointo & Paéz/ Cord. Central, E ofSantander de Quilichao, SE ofToribio, NE Planadas & Río blanca ofPmo. de Las Moras, SW comer ofParque Nacional Natural Nevado del Huila, S ofthe Nevado
05°37'N
73°08'W
Boyacá
Tora& Tuta
Cord. Oriental, E ofTunja, E ofToca, W ofPesca
07"16'N
72°53'W
Santander
Tona
Cord. Oriental, SE of Pmo. Rico and connected with it, 6 km NE ofBucaramanga, E ofBerlín
3500
05°37'N
72°52'W
Boyacá
Aquitania (Puebloviejo)
Cord. Oriental, NE ofLaguna de Tota, between Somagoso and Vado Hondo, headwaters ofRío Cusiana
3000
06°07'N
72°48'W
Boyacá
Cord. Oriental, above Córdoba, near Susacón
Cord. Oriental, E ofCord. de Consuelo, NE ofTutasá
s;: s;: o
(1j
¡;a [/)
o
"rj
>-l
::e (1j
z
(1j
::E
o-< ~ ttl o>-l >z ñ
>l' o >E3 (1j
z
<o ~
00
~
N ame
Elev. (m)
Latitude
Longitude Division 1
Division 2
Savelillo, Pmo. de
Notes Cord. Central, N of Pmo. de Huila, due W of Neiva, SE of Jagual, S ofPmo. de Iraca
Seca, Pmo. de Laguna
3630
05°02'N
73°55'W
Cundinamarca
Tausa
Cord. Oriental, between Cagua and San Cayetano, E ofZipaquirá
Siachoque, Pmo. de
3500
05°27'N
73°ll'W
Boyacá
Siachoque, Viracachá & Rondón
Cord. Oriental, ESE of Tunja, W of Laguna de Tota, E of Ramiriquí, NW ofPmo. de las Alfombras and connected with it; also called Alto Siachoque
Sibaté, Pmo. de
3570
Cundinamarca
Siberia, Pmo. de (La)
3500
Cundinamarca
La Galera
Cord. Oriental
Siecha, Laguna de
3600
04°46'N
72°48'W
Cord. Oriental, ca. 23 km SE ofBogotá
Cundinamarca
Cord. Oriental, in páramo
Siejo, Pmo.
See Cerro El Viejo
Simón Bolívar, Pico
5775
10°50'N
73°42'W
Magdalena
Sierra Nevada de Santa Marta, NW ofValledupar, between municipalities of Aracataca and Santa Marta
Simons, Pico (El)
5660
!0°5l'N
73°43'W
Magdalena
Sierra Nevada de Santa Marta, NW ofValledupar, between municipalities of Aracataca and Santa Marta
Siquitán, Pmo. de Sirpes, Pmo.
See Pmo. Chimbalán 2200
Siscunsí, Pmo. de
3700
Sisga, Pmo. El
2850
Antioquia 05°22'N
72°50'W
Boyacá
San Rafael & Guatapé Sogamoso
;¡,.
Cord. Oriental, may be part ofPmo. de Toquilla
;¡,.
Cord. Oriental, near Chocontá
Socabón, Pmo. del
Cauca
Cord. Central
Socha, Pmos. de
Boyacá (?)
Cord. Oriental, Sierra Nevada del Cocuy
Soledad, Pmo. La(!)
3600
Soledad, Pmo. La (2)
3600
Sonsón, Pmo. de
3340
03°55'N 05°43'N
74°49'W 75°!5'W
Sora, Pmo. de
Tolima!Valle
"O
Cord. Central; not true páramo
Cundinamarca
;;<!
3::
o
(ll
Cord. Central, E of Santa Lucía, SE ofBarragán
Huila
San Agustín
This may be Cauca, Mpio. Santa Rosa, 01 o40'N, 76°34'W
Antioquia
Sonsón, Argelia & Nariño
Cord. Central, N ofPmo. Arboleda, 65 km SE ofMedellín, NE of Aguadas, E of Sonsón; an isolated peak with subpáramo vegetation at top
Boyacá
Sora & Cucaita
Cord. Oriental, surrounds Laguna de Suesca; see Hettner, 1892; and Guhl, 1966: 62, 110 Macizo Colombiano; incl. Volcán Sotará (4850 m), S ofTimbío
Sotará, Pmo. de
4100
02°!2'N
76°3l'W
Cauca
Sotará (Paispamba) & La Vega
Soté, Pmo. de
3200
05°39'N
73°!9'W
Boyacá
Tunja & Cómbita
Sulibata, Pmo. de
5375
!0°40'N
73°30'W
Magdalena
Suma Paz, Pmo. de la
::0
\0
~
Cord. Oriental, páramos N and S of Soté, 15 km N ofTunja Sierra Nevada de Santa Marta See Pmo. de Sumapaz
:;: --._)
~
Elev. (m)
Latitude
Longitude Division 1
Sumapaz, Pmo. de
4285
03°45'N
74°25'W
Cundinamarca/Meta/ Huila
Surivaca, Pmo. de
4000
l0°53'N
73°27'W
Cesar/La Guajira
Riohacha
Boyacá
Aquitania (Puebloviejo)
N ame
Division 2
Santander
Sutamarchán, Pmo. de
Cord. Oriental
~ m
Cord. Oriental, N ofVetas
o
See Pmo. de la Pastora
Tábano, Pmo. de Tábano, Pmo. de(!)
3300
01 °10'N
77o12'W
Putumayo/Nariño
Tablazo, Pmo. (de El)
3500
04°52'N
74°13'W
Cundinamarca
05°28'N
75°23'W
Caldas
Tacará, Pmo.
3000
Tacines, Pmo. de
3500
Nariño
15 km E ofPasto toward El Encano Subachoque
Cord. Oriental, ca. 43 km NNE ofBogotá, W ofSubachoque Cord. Central, E ofSalamina, SE ofPacora, NW ofPmo. San Félix and connected with it; may not be páramo
Pasto
Serranía del Morasurco
Tairona, Pico
5000
!0°47'N
73°39'W
Magdalena
Aracataca
Sierra Nevada de Santa Marta
Tajumbina, Pmo. de
3600
01 °24'N
76°47'W
Nariño/Cauca
El Tablón & La Cruz
Cord. Central, NE of Pasto, S ofLa Cruz, slopes ofCerro Tajumbina and Cerro Potosí; incl. also parts ofVolcán Doña Juana and Cerro Animas
Tamá, Pmo. de
3329
07"25'N
72°26'W
N Santander
Toledo
Cord. Oriental, E ofPmo. de Tierra Negra and connected with it, SW ofCúcuta, E ofPamplona, shared with Venezuela
Tamaná, Cerro
4200
05°02'N
76°JJ'W
Risaralda/Chocó
Tambo, Pmo. del
3300
01 °0l'N
77°5l'W
Nariño
Macizo Colombiano, Km 16 along Cumbal-Miraflores rd.
Tambor, Pmo. El
3800
04°02'N
75°45'W
Tolima!Valle
Cord. Central, S ofPmo. Barragán, S ofGenova, E ofBarragán
Cord. Occidental, W ofSantuario, S ofPueblorrico
Taquina, Pmo.
2800
La Guajira
Riohacha
Tasajeras, Pmo. (!)
3800
07"27'N
72°55'W
Santander
Suratá
Tasajeras, Pmo. (2)
3800
04°03'N
74°16'W
Cundinamarca
05°00'N
76°05'W
Risaralda/Chocó
Tatamá, Cerro
3950
Tausa, Pmo.
3200
Tauso, Pmo. El
3500
01 °04'N
77°20'W
~
~
"' o""rj >-l
:r: m zm
:;e o--< ?::1
;>:: tJ:j
o>-l > z ()
> t-< Cl
>
§
Putumayo
Tambillo, Pmo. de
00
Sierra Nevada de Santa Marta; also called Pmo. de Sulibata or spelled Suliveta or Surlibáka; equals Pmo. Sulibata from Magdalena See Pmo. de Surivaca
3600
0\
Cord. Oriental, SE ofVi!larica, 25 km S ofBogotá; incl. Pmos. de Mundo Nuevo, de Las Animas, and de Nevado (4560 m) and the Altos de Andabobos, Rabona, Tortica (or Fraile) and Ramírez; also called Cerro Nevado de Sumapaz and Macizo de Sumapaz
Surlibáka, Pmo. de Suse, Pmo. (de)
Notes
~
Sierra Nevada de Santa Marta; not páramo Cord. Oriental, SE of Cachiri Cord. Oriental, S ofFusagasugá, S ofPmo. Andabobos, part of the larger Pmo. de Sumapaz Cord. Occidental, SW ofSantuario, NW of Ansermanuevo
Cundinamarca
Tausa
Cord. Oriental; a village, not páramo
Nariño
Tan gua
Cord. Centro-Oriental, SE ofPasto along rd. toLos Alisales
<o r-'
00
.¡,..
N ame Umbita, Pmo.
Elev. (m)
Latitud e
Longitud e Division 1
3000
05°13'N
73°29'W
Division2
Antioquia
Cord. Central, possibly vicinity of Sonsón
Urrao, Pmo. de
3500
Usaquén, Pmo. de
3060
Cundinamarca
Usme, Pmo.
3200
Cundinamarca
Urrao
Cord. Oriental, 18 km S ofBogotá, SE ofUsme
Cord. Oriental, Macizo de Bogotá
Utanquer, Pmo.
3300
Nariño
Sapuyes
Nudo de Los Pastos
Uvita, Pmo. de La
4000
06° 18'N
72°3l'W
Boyacá
La Uvita
Cord. Oriental, E of Soatá, W ofPmo. de Rechíniga and connected with it, SW of Sierra Nevada del Cocuy
Vado Ancho, Pmo. de
3700
06°55'N
72°40'W
Santander
Cerrito
Cord. Oriental, S ofPmo. del Almorzadero
Boyacá
Cord. Oriental, E slopes, in the valley ofRío de Cusiana
Valencia, Pmo. de
4200
01 °55'N
76°32'W
Cauca!Huila
N ofPmo. de Las Papas (or is it the same locality?); elevation and coordinates approximate
Valle Bonita, Pmo. de
3800
03 °45'N
76°00'W
Valle
Cord. Central, NE ofPalmira, E ofPmo. de Los Domínguez
Valle Lagunillas, Pmo. de
4060
Boyacá
Cord. Oriental, Sierra Nevada del Cocuy
Valledupar, Sierra de
3600
l0°35'N
72°48'W
o
Cord. Occidental
Antioquia
Vadohondo,Pmo.de
...¡
Cord. Oriental, NE ofChocontá, E ofVillapinzón
Cundinamarca!Boyacá
Unión, Pmo. de La
Notes
Magdalena (CO)/Zulia (VE)
. Cord. de Perijá, E of Villanueva, along Colombia/Y enezuela border Cord. Central, 45-50 km SE of Cajamarca, close to Río Anaime; also ·called Pmo. Los Valle
~
tri
~
o ?a Cll o "rl
..., ::r: tri
ztri
::E
Vegas, Pmo. de Las(!)
3800
Santander
Cord. Oriental
Vegas, Pmo. de Las (2)
3800
Valle
Cord. Central, above Tuluá and Bugalagrande
o--< ~ IJ:I ...,o >z ñ >l'
Risaralda
Cord. Central; bryophyte locality for Breutelía bríttoniae (Daniel & Ramón Guevara 6297)
§
Cord. Oriental
z
Valles, Pmo. de Los
3800
04°l5'N
75°3l'W
Tolima
Vasto, Pmo. del
3800
06°02'N
72°58'W
Boyacá
Cerinza
Santander
Suratá
Vega de Páramo
Vera Cruz, Pmo. de Verde, Pmo.
3100
06°06'N
72°34'W
Boyacá
Jericó
Cord. Oriental, NW ofBelén Cord. Oriental; a group ofhomes, nota páramo
Verde, Pmo. El
4200
06°22'N
72°22'W
Boyacá
Cord. Oriental
Verde, Pmo. de Laguna (1)
4200
06°34'N
72°24'W
Boyacá
Cord. Oriental, NE ofEl Cocuy, E ofLas Mercedes
Verde, Pmo. de Laguna (2)
3700
05°13'N
74°00'W
Cundinamarca
Cord. Oriental, between Cogua and San Cayetano, ca. 60 km NNW ofBogotá
Vergara, Pmo. (de)
3400
05°37'N
73°06'W
Boyacá
Ve¡jón, Pmo. del
3600
04°32'N
74°04'W
Cundinamarca
Cord. Oriental, Macizo de Bogotá, side nearest Choachi, SE of Bogotá
Vetas, Pmo. de (Las)
2800
Santander
Cord. Oriental; type locality of Espeletia santanderensis
Tuta
Cl
>-
tri
Cord. Oriental, NE ofTunja, NW ofLaguna de Tota
<o l' 00
-~'"
~
N ame Viejo, Cerro el
Elev. (m)
Latitude
Longitude Division 1
4100
or27'N
72°54'W
Division2
N Santander/Santander
Cord. Oriental, SW ofCúcuta, NE ofBucaramanga; may be the same as Pmo. Viejo, which has been misspelled as Pmo. Siejo
'O 'O
~
See Cerro El Viejo
Viejo, Pmo.
See Pmo. Bijagual (de)
Vijagual, Pmo. de Villapinzón, Pmo. de
Cundinamarca
Cord. Oriental; may be spelled Villa Pinzón
Visitador, Pmo. del
Cord. Central
Yacuanquer, Pmo.
3400
Yechiquín, Cerro
3650
Yerbabuena, Pmo. de
3150
Yula, Pmo. de
3400
Yunguilla, Pmo de Zea, Cerro La
3800
Zipaquirá, Pmo. de
3200
Zumbador, Pmo.
Notes
Nariño
Ospina
Nudo de Los Pastos
Magdalena
Aracataca
Sierra Nevada de Santa Marta
Tolima
Ronces valles
Cord. Central
04°57'N
73°44'W
Cundinamarca
01 °54'N
76°42'W
Cauca
San Sabastián
Cord. Central; also spelled Junguilla
Cauca
San Sabastián
Macizo Colombiano
05°02'N
74°00'W
Cundinamarca
Cord. Oriental, between Zipaquirá and Pacho
or57'N
72°45'W
-N Santander
Cord. Oriental
Cord. Oriental
"O
> ~ 3::: o (/)
-..)
172
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
[VOL. 84
Venezuela James L. Luteyn and Mauricio R. Gavilanes A.
The Cordillera Oriental ofColombia splits into two unequal spurs as it enters Venezuela: The smaller one goes directly north and is called the Sierra de Perijá, and the larger one goes in a northeast direction and is generally called the Cordillera de Mérida. The Cordillera de Mérida, in turn, transects the Venezuelan Andes in a southwest to northeast direction, ranging over 450 km, from the Táchira depression near San Cristóbal to the Barquisimeto~Carora depression in southwestern Lara. The páramos of Venezuela are centered in the states ofMérida, Táchira, and Trujillo, with smaller numbers in Barinas, Lara, Apure, and Portuguesa. The primary mountain ranges in the state ofMérida aré the Sierra del Norte (or also called the Sierra de La Culata), which is north of the city of Mérida, the Sierra Nevada de Mérida to the south of Mérida that includes the Parque Nacional Sierra Nevada, and the Sierra de Santo Domingo, northeast of Mérida. The highest point in Venezuela is Pico Bolívar (Mérida state) at 5007 m. In the state of Trujillo, the eastern continuation ofthe Andes mountain range is called the Cordillera de Trujillo, which has severa! branches such as the Ramal de Calderas that continues into the state of Barinas, Ramal del Rosario east-northeast of Boconó, and the Sierra Morena de Trujillo that continues into the state of Lara. The primary ranges in Táchira are the Sierra del Batallón, Sierra La Cimarronera, Sierra de Callejón Colorado, and Sierra de Palo Grande. In the state of
Zulia, along the Colombian border, the extreme northern limit of the Cordillera Oriental of Colombia is known as the Sierra de Perijá, with its southern part krú:íwn as the Serranía de Los Motilones and the northetn part the Serranía de Valledupar. Jahn (1931) estimated that páramo occupied 4100 km 2 in Venezuela. According to many herbarium labeis, the Venezuelan páramos often seem to begin at lower elevations than those ofthe other countries, e.g., 2500~2800 m. This may be true in sorne rare cases since in general the páramos ofVenezuela are somewhat drier than those of Colombia and northern Ecuador. It is my (JLL) experience, however, that most ofthese areas are usually not páramo; rather, they are areas that have been severely disturbed by man and are now almost totally dominated by the fern Pteridium aquilinum, with scattered Monochaetum (Melastomataceae) and Dodonea viscosa (Sapindaceae). Ido not in this book consider these areas páramo (but see also Bono, 1996). Venezuelan national párks that include páramo are Sierra Nevada, Sierra de la Culata, Dinira, El Tamá, Perijá, Guaramacal, and Páramo Batallón~La Negra (cf. Gabaldón, 1992). For additional discussion ofVenezuelan páramo vegetation, see Bono, 1996; Fariñas & Monasterio, 1980; Goebel, 1891; Hitchcock, 1954; Jahn, 1931; Monasterio, various papers cited; Tamayo, 1958, 1975; Tillett, 1978; Vareschi, 1970; and Viloria P., 1993.
'D 'D
Elev. (m)
Latitude
Longitude
Division 1
Division2
Notes
3900
08°22'N
71 o 14'W
Mérida
CampoElías & Arzobispo Chacón
Sierra Nevada de Mérida, 20 km SW ofMérida, Pueblos del Sur area, SE of Las Acequias, N of Pmo. Aricagua and connected with it; also ca1led Pmo. de Las Lajas or Pmo. de La Laja
Adobe, Pmo. El
3265
08°0l'N
72°00'W
Táchira
Cárdenas & Jáuregui
SE ofEI Cobre
Agrias, Pmo. Las
3362
or59'N
72°05'W
Táchira
Jáuregui/Sucre
S ofEl Cobre, NW ofQueniquea
Agua Blanca, Pmo. de
4600
08°48'N
71 °02'W
Mérida
Libertador, Justo Briceño & Andrés Be11o
Sierra de la Culata, 20 km NNE ofMérida, E ofPmo. Tucaní and connected with it, N ofPmo. El Escorial and connected with it
Agua Fría, Pmo. de
3121
09°!6'N
70°08'W
Trujillo
Boconó
Ramal del Rosario, ENE of Boconó, E ofMosquey, SW of Pmo. de Los Rosarios and connected with it; is it páramo?
Agua Linda, Pmo. de
1150?
10°ll'N
70o29'W
Lara
Torres
Agua de Obispo, Pmo. de
2500
Aguada, Pmo. Estación La
3450
08°34'N
AguiJa, Pmo. del
4126
Alcobé, Pmo. (El)
N ame Acequias, Pmo. de
See Pmo. Apure
Acure, Pmo.
Doubtfu11y páramo
Trujillo
Notpáramo
71 °06'W
Mérida
Sierra Nevada de Mérida, Parque Nacional Sierra Nevada, station along the Mérida Teleférico
08°5l'N
70°50'W
Mérida
Rangel & Miranda
NE ofMérida, 18 km SW ofTimotes, vicinity ofPico El AguiJa (highest part ofPmo. Mucuchíes); also ca1led Pmo. Pico AguiJa
3265
08°0l'N
72°05'W
Táchira
Grita & Uribante
SE ofEl Cobre
Almorzadero, Pmo. (1)
2843
ors9'N
72°06'W
Táchira
Jáuregui
Sierra de Palo Grande, dividing line between Ríos Torbes and Lobatera, E ofMichelena; also ca11ed Pmo. Zumbador
Almorzadero, Pmo. (2)
1598
Táchira
Ramal del Uribante; not true páramo
Almorzadero, Pmo. (3)
1378
Yaracuy
Macizo de Nirgua; not true páramo
Almorzadero, Pmo. de (4)
4200
Mérida
Altamira, Pmo. de
3320
08°03'N
71 °33'W
Mérida
Alto (de) Timotes
.., ~
~ 2'::: o IZl
NE ofMérida anda little NE ofPmo. Piedras Blancas area Rivas Dávila, Arzobispo 22 km E of Pregonero, N of Pmo. El Pabellón; incl. Pico de Chacón & Uribante Horma See Pmo. de Timotes
Alto El Morro
See Pmo. El Morro
Alto Páramo de La Negra
See Pmo. (de) La Negra
Alto de Arenales
See Pmo. Arenales
Alto de Arenas
See Pmo. Arenales
Alto de Tomón
See Pmo. de Tomón See Pico El Oso
Alto del Oso Alto del Páramo
.:S
1530
Monagas
Macizo de Caripe; not páramo
:; w
N ame
Elev. (m)
Latitud e
Longitude
Division 1
Division 2
Alto del Páramo del Morro
...., ~
See Pmo. El Morro
Angaraveca, Pmo.
2793
Táchira
Angaraveca, Pmo. de
2793
Táchira
Angustias, Pmo. Las
2350
Apartaderos, Pmo. (de)
3550
Aranguren, Pmo. (de, Los)
3000+
Árbol Redondo, Pmo.
2480
09°23'N
70°18'W
Arenales, Pmo.
3810
09°02'N
70°36'W
Aricagua, Pmo. de
3550
08°l5'N
71 °13'W
08°06'N
71 °27'W
Ayacucho Not true páramo
Mérida
Arzobispo Chacón
SSE ofLibertador (Canaguá)
Mérida
Miranda or Rangel
Sierra de Santo Domingo
Mérida
Asucati, Pmo. de Atajo, Pmo. El
Notes
Sierra de Santo Domingo (see López-Zent, 1993)
Trujillo
Trujillo & Boconó
Cord. de Trujillo, SW ofBurbusay; not true páramo
Mérida/Trujillo
Miranda!Urdaneta
Cord. de Trujillo; also called Alto (de) Arenas and Pico Arenas
Mérida
Libertador & Arzobispo Chacón
N of Aricagua, 41 km S ofMérida, S ofPmo. Buenos Aires, SE ofPmo. Don Pedro, 10 km E ofMocomboco
Trujillo & Boconó
WNW ofBoconó, N ofPmo. Los Pozos, SW ofPmo. La Cristalina
Jiménez & Morán
Sierra de Portuguesa, E ofGuarico; not true páramo
Mérida 3200
09°l6'N
70°22'W
Trujillo
Baho, Pmo. El
2800
08°52'N
70°4l'W
Mérida
Bajó, Pmo.
2176
09°38'N
69°4l'W
Lara
Balcón, Pmo. El
4200
Banco, Pmo. El
4200
08°48'N
70°57'W
Mérida
Rangel, Libertador & Justo Briceño
Sierra Neyada de Mérida, NE ofMérida
Bandera, Pmo.
3000?
or28'N
72°25'W
Táchira
Junín
N ofPmo. de Tamá and connected with it, due E ofVilla Páez
SW of Santo Domingo; not true páramo
Mérida
Barbacoas, Pmo. de
· Sierra de Santo Domingo?
Trujillo/Barinas
Barrial, Pmo. El
a::trJ a:: o
~ (/)
o'T1 ..., ::r: trJ
ztrJ
::E
o-< ~ td o...,
;¡,.
08°05'N
71 o33'W
Mérida
Arzobispo Chacón
Ramal de Uribante, SW ofLibertad (Canaguá)
Bartola, Pmo.
3492
09°0l'N
70°28'W
Trujillo/Barinas
Boconó/Bolívar
Ramal de Calderas, SSW of Niquitao village, N of Calderas, NE of Pmo. Motumbo and connected with it, W of Pmo. de Rojas and connected with it, ESE ofPmo. Calderas
Batallón, Pmo. (de, del El, del, El)
3507
08°05'N
71 °55'W
Táchira
Jáuregui & Uribante
Sierra del Batallón, NW ofPregonero, 7 km E ofLa Grita, 15 km SW ofBailadores, S ofPmo. Rosario and connected with it, N ofPmo. Sumusica and connected with it
Biriguaca, Pmo.
3421
Mérida
Blanca, Pmo. La
3810
Táchira
Blanco, Pmo. de El
3450
Mérida
Misspelling for Pmo. El Banco?
Boconó, Pmo.
2600
Trujillo
Probably a general name for páramos around Boconó, but seems too low for true páramo
z
ñ ;¡,. r
~ ~ trJ
z
Sierra La Cimarronera, S of Pmo. La Ciénega and connected with it
<o r-'
00
~
N ame
Elev. (m)
Latitud e
Longitude
Division 1
Division2
Notes
Bolívar, Pico
5007
08°33'N
71 °03'W
Mérida
Libertador
Sierra Nevada de Mérida (highest point), 10 km SE ofMérida; also called Pmo. La Columna; páramo surrounds it Sierra Nevada de Mérida
Bonpland. Pico
4883
08°33'N
71 aoo·w
Mérida
Libertador
Boquerón, Pmo.
2600
08°04'N
71 °28'W
Mérida
Arzobispo Chacón
E ofPmo. Altamira, SW ofPmo. Botalón; true páramo?
Botalón, Pmo.
2600
08°05'N
71 °25'W
Mérida
Arzobispo Chacón
NE ofPmo. Boquerón and connected with it; true páramo?
08°19'N
7lo1l'W
Mérida
Campo Elías, Arzobispo N ofPmo. de Aricagua and connected with it Chacón & Libertador
08°5l'N
70°54'W
Mérida
Rangel & Justo Briceño
08°47'N
70°51'W
Mérida
09°07'N
70°26'W
Trujillo
Boconó & Urdaneta
Trujillo
Urdaneta & Boconó
09°03'N
70°25'W
Trujillo/Barinas
Boconó/Bolívar & Obispos
Ramal de Calderas, S of Niquitao, W of Masparrito, ENE of Pmo. Bartolo and connected with it, N of Pmo. de Rojas and connected with it, SW ofPmo. Ortiz and connected with it Sierra de Callejón Colorado, WNW ofQueniquea
Buenos Aires, Pmo. de Buitre, Pmo. El
4400
Cabimbú, Pmo. de
4000
Cachaco, Pmo. de
3700
Calderas, Pmo. de (las)
3642
Callejón Colorado, Pmo. (de)
3350
Camello, Pmo. Campanario, Pmo. (El)
4325
Cañada, Pmo. de La
2700 2740
Canaveral, Pmo. El Caracoles, Pmo. de Los
4736
Carbonera, Pmo. La (1)
4000
Carbonera, Pmo. La (2) Cardenillo, Pmo. El Caricuena, Pico
Part ofPmo. de Mucuchíes, SE ofChachopo; also called Pmo. Los Buitres See Pmo. El Buitre
Buitres, Pmo. Los Caballo, Pmo. Mesa del
Canaguá, Pmo. (de)
\0 \0
.:3
4480
Also called Mesa de Los Pozos W of Niquitao, lower slopes on E side ofTeta de Niquitao, below Pmo. La Vieja
07"56'N
72°04'W
Táchira
Jáuregui & Cárdenas
07"27'N
72°ll'W
Táchira/COLOMBIA
San Cristóbal/N Santander
08°40'N
71 °!6'W
Mérida
Campo Elías & Andrés Bello
Sierra de La Culata, NW of Mérida, incl. Pico Campanario (4299 m), W ofPmo. La Pedregosa and connected with it, SW ofPmo. Los Conejos and connected with it
Trujillo
Boconó
Part ofPmo. de La Cristalina
08°13'N
71 °30'W
Mérida
Arzobispo Chacón
Close to Pmo. Pozo Negro and Pmo. El Cupís, S ofPmo. Las Nieves; not true páramo; also called Latalito; may equal Pmo. El Molino (1) (B. Briceño, pers. comm.)
Mérida
El Aracay-La Quebrada del Padre area
Mérida
Cord. del Norte (or de La Culata); incl. Pico Caracoles (4736 m)
Mérida
Sierra de Santo Domingo
Mérida
Cord. del Norte (or de La Culata), Mérida-La Azulita rd., above town ofLa Carbonera; true páramo?
Mérida
Sierra Nevada de Mérida
>;j
> ~ ~ o<Zl
See Pmo. Caricuena --.¡ Vl
Name Caricuena, Pmo.
Elev. (m)
Latitud e
Longitude
Division 1
Division 2
Notes
3000
or53'N
71 °50'W
Táchira
Cárdenas & Jáuregui
ESE ofSan José de Bolívar, NW ofLa Fundación; also called Pico Caricuena
Carimbú, Pmo.
3000
Trujillo
Urdaneta
See Pmo. de La Teta de Niquitao
Carreros, Pmo. de Los
3224
08°13'N
71 °47'W
Mérida
Rivas Dávi1a
ESE ofBailadores, N ofPmo. Las Tapias and connectd with it, N of Pmo. San Pedro and connected with it, SW of Pmo. Guaraque and connectd with it; is this part ofPmo. Las Tapias?
Castillejo, Pmo.
3558
09°03'N
70°23'W
Trujillo/Barinas
Boconó/Obispos
Ramal de Calderas, S of Niquitao, W of Pmo. Masparro and connected with it, NE ofPmo. Ortiz and connected with it
Cedro, Pmo. El
3600
08°04'N
71 °53'W
Táchira
Uribante
NW ofPregonero
Cendé, Pmo. (de)
3583
09°34'N
70°07'W
Trujillo/Lara
Boconó & Carache/Morán
Sierra Morena de Trujillo, NW, N and NE slopes of Pico Cendé, S ofPmo. Jabón, W ofEl Jabón, 40 km NE ofTrujillo, SW ofHumocaro Alto; sorne place Pico Jabón to the N ofPmo. (de) Cendé and Pmo. Jabón toS ofPmo. (de) Cendé
Chacantá, Pmo. de
Mérida
2000
Not true páramo See Pmo. de Chachopo
Chachapos, Pmo. de Chachopo, Pmo. de
3200
08°56'N
71°46'W
Mérida
Miranda
Part ofPmo, de Timotes; also called Pmo. de Chachapos
Chegué, Pmo.
3894
09°04'N
70°49'W
Mérida!Trujillo
Miranda & Justo Briceño/Escuque
NE ofPiñango, N ofPmo. Malpica; also called Pmo. Cheque
Cheque, Pmo.
See Pmo. CIÍ.egué
China, Pmo. La
See Pmo. (de) SanTelmo Sierra Nevada de Mérida, Pueblos del Sur area
Mérida
Chirivital, Pmo. de Chono, Pmo.
2800
Chorote, Pico
3452
Chorrerón, Pmo. de
2750
Chorro Blanco, Pmo.
3680
Chorro, Pmo. de El (del)
2800
09°0I'N
70°32'W
Mérida
Rangel
Mérida/Trujillo
Miranda/Boconó & Urdaneta
09°06'N
70°3I'W
Trujillo
0\
s;: til s;: o ¡;:;
en
o
"rj
>-j
::r: til
ztil
:E o--<
::0 :;<: tD
o>-j z> ñ > r-" Cl
>
E3
Mérida
Chorrotes, Pmo. de los
til
Urdaneta & Boconó
18 km S ofSantiago de Trujillo, 15 km E of Jajó, W slopes of Teta de Niquitao, E ofPmo. del Tostado and connected with it
Mérida
Near Mucurubá
Trujillo
NE of Jajó; possibly the same as Pico Chorote
Ciénega, Pico La Ciénega, Pmo. La
Near Mucurubá, near Hacienda de Cacúte; páramo?
.....,
z
See Pmo. La Ciénega 3868
or56'N
71 °59'W
Táchira
Jáuregui & Uribante
Sierra La Cimarronera, part ofPmo. La Cimarronera, N of San José de Bolívar; also called Pico La Ciénega
<o ¡-'
00
.¡o,.
Elev. (m)
Latitude
Longitude
Division 1
Division2
Notes
3795
or5s'N
71 °55'W
Táchira
Uribante, Jáuregui & Cárdenas
15 km SSE ofLa Grita, 16 km W ofPregonero, NNE ofSan José de Bolívar, E ofPmo. El Rosal and connected with it, S of Pmo. Sumusica and connected with it; incl. Pmo. La Ciénega, Pmo. El Molino, and Pmo. El Lajón (1)
Cobre, Pmo. (El)
3675
or2s'N
Táchira
Junín
E ofPmo. de Tamá; Pico El Cobre located at 3613 m
Cocuy, Pmo. El
4400
08°30'N
7r22'w 71 o07'W
Mérida
Libertador
Sierra Nevada de Mérida, S of Mérida; also caiied Pmo. El Paramito and equal to Pmo. del Morro (B. Briceño, pers. comm.)
Colorada, Pmo. La
3624
Táchira
Jáuregui
Coloradas, Pmo. (de) Las
3635
08°19'N
71 °20'W
Mérida
Campo Elías & Arzobispo Chacón
SW of Mérida, SW of Pmo. San José and connected with it; incl. Pico El Rejo (3635 m)
Colorado, Pmo. (de) El
3300
or55'N
72°05'W
Táchira
J áuregui/Sucre
E ofQueniquea, connected with Pmo. El Zumbador and Pmo. El Portachuelo; also cailed Pmo. de Los Colorados and sometimes considered a sector ofPmo. Zumbador
N ame Cimarronera, Pmo. La
'O 'O
c::s
See Pmo. (El) Cobre
Cobre, Pico El
See Pmo. (de) El Colorado
Colorados, Pmo. de Los
See Pico Bolívar
"O
Concha, Pico La
4922
08°34'N
71 o03'W
Mérida
Libertador
Sierra Nevada de Mérida, W ofMérida, N ofPico Bolívar; also caiied Pico La Garza
S:;;::
Conejos, Pmo. (de) Los
4000
08°44'N
71 °!3'W
Mérida
Andrés Belio, Campo Elías & Libertador
Sierra de La Culata, 15 km NW ofMérida, near San Isidro, N ofPmo. Campanario and connected with it
Corazón, Pmo. El
3000
09°12'N
70°24'W
Trujiiio
Trujiiio & Boconó
W ofTostos, WSW ofBoconó, N ofNiquitao, S ofPmo. Los Pozos and connected with it
Corcovada, Pmo. de La
3650
08°07'N
70°44'W
Mérida
Rangel
Sierra de Santo Domingo, SE of Apartaderos; incl. lagunas Mucubají and La Negra
3000
09°l9'N
70°20'W
Trujillo
Trujiiio & Boconó
Cord. de Trujillo, SE ofTrujillo, NW ofBoconó, NE ofPmo. El Atajo
or25'N
72°25'W
Táchira
Cruces, Pmo. (de) Las
4200
08°52'N
70°52'W
Mérida
Cuchicuchi, Pmo.
2147
Táchira
Cucuchica, Pmo.
3234
Mérida
Cueva Negra, Pmo. de
2400
Columna, Pmo. La
>
o
IZl
Cordero, Pmo. de Los Cristalina, Pmo. (de) La Cristo, Pmo. de
08°10'N
71 °38'W
Mérida
Colombia/Venezuela border, close to Pmo. de Tamá Rangel
W ofMucuchíes, N ofSan Isidro, forms the SE end ofPmo. de Mucuchíes Not true páramo
Rivas Dávila & Arzobispo Chacón
23 km SE ofTovar, 23 km NE ofPregonero, 7 km N ofMesa de Quintero; not true páramo
::::; -.)
N ame
Elev. (m)
Latitud e
Longitude
Division 1
Division 2
Notes
Culata, Pmo. (de La)
4530
08°50'N
71 °03'W
Mérida
Libertador & Andrés Bello
Cord. de La Culata, 15 km N ofMérida, N ofTabay, W and S slopes ofCerro La Torre, SW ofPmo. Tucaní and connected with it, NE ofPmo. Los Leones and connected with it, ENE ofPmo. Los Conejos and connected with it [incl. Pico La Culata (4620 m)]
Cupís, Pmo. (de El, El)
3200
Mérida
Arzobispo Chacón
Between San José and Veguilla (La Be guilla), part ofthe larger Pmo. de San José, close to Pmo. Canaguá and Pmo. Pozo Negro
Curas, Pmo. Los
4400
08°42'N
71 °JO'W
-...)
00
Mérida See Pmo. Guirigay
Curiguay, Pmo. Destierro, Pmo. del Don Alfonso, Pmo.
3600
Don Pedro, Pmo. (de) (1)
3790
08°2l'N
71 °04'W
Don Pedro, Pmo. (2) Duque, Pmo. de (El)
3260
Durí, Pmo. de
3000
09°02'N
70°41'W
Escorial, Pmo. (El, del)
3500
08°46'N
71 °06'W
Esdorá, Pmo.
3245
Espejo, Pmo.
4550
Estrella, Pmo. (La)
3798
Fortuna, Pmo. La
2230
Fría, Pmo. (La)
4000
Mérida
Sierra del Norte, near Campanario
Mérida
Sierra Nevada de Mérida
Mérida!Barinas
Libertador/Pedraza
Mérida
Justo Briceño & Miranda Sierra de La Culata
Táchira
Jáuregui
La Grita-Pueblo Hondo rd., ca. 5 km S ofEl Cedrillo
Trujillo/Mérida
Urdaneta!Miranda
N ofPueblo Llano
Mérida
Rangel & Libertador
E ofSierra.de La Culata, ENE ofMérida Between Alto ~de Pampanito and Pmo. de Tuñame, NE of Val era
Trujillo 08°32'N
08°57'N
08°36'N
71 °03'W
70°41'W
71 °02'W
Mérida
Mérida/Trujillo
Sierra Nevada de Mérida, Pueblos del Sur area, NNE of Aricagua, SSE ofMérida, 20 km SE ofEl Morro
Libertador
Miranda/?
Sierra Nevada de Mérida, Parque Nacional Sierra Nevada, SW of Pico Bolívar, station along the Teleférico de Mérida; incl. Laguna El Espejo at 4000 m and Pico Espejo at 4550 m Cord. de Trujillo, NE of Chachopo, SE of Timotes, S of La Mesa de Esnujaque (Trujillo)
Táchira
Not true páramo
Mérida
N part of Sierra Nevada, SE ofTabay; incl. Laguna La Fría at 3000 m; also called Pmo. de Frías; elevation doubtful, most likely lower
Frías, Pmo. de
3:: tTJ 3::
o ;;:; IZl o.., ....J ~
tTJ
ztTJ
::E --<: o
~
ttl
o
....J
:>
z
ñ
:> t-'
o :> ES tTJ
z
See Pmo. (La) Fría
Gallo, Sierra del
See Pmo. Los Nevados
Garza, Pico La
See Pico La Concha
Gavidia, Pmo.
3800
08°38'N
70°58'W
Mérida
Libertador
Sierra Nevada de Mérida, separation point with Sierra de Santo Domingo; elevation not certain, most likely lower
Gavilán, Pmo. (del)
4000
08°5l'N
70°48'W
Mérida
Libertador & Miranda
E ofPmo. de Mucuchíes, NE of Apartaderos; Pico Gavilán rises to4248 m
Gaviria, Pmo.
4402
Mérida
Sierra Nevada de Mérida to Santo Domingo
<o r
00
~
N ame Gonzáles, Pmo. de Las
Elev. (m)
Latitud e
Longitude
4300
Division 1
Division 2
Mérida
Notes Sierra del Norte, headwaters o fRío La Gonzáles, community of Valle del Campanario is located here; also spelled González
Gordo, Cerro
See Pmo. (del, El) Salvaje
Granates, Pmo. (de Los)
3900
08°47'N
70°4l'W
Mérida/Barinas
Rangel/Bolívar
Sierra de Santo Domingo, S of Santo Domingo, between El Hato and Alto del Morato; also spelled Granate
Guache, Pmo. (de, El)
3060
09°38'N
70°07'W
Trujillo/Lara
Morán & Carache/?
Cord. de Trujillo (N part), E ofCarache, N ofPmo. Las Rosas
08°02'N
71 °32'W
Mérida
Arzobispo Chacón
Cord. de Mérida, N of Guaimaral
3666
08°05'N
71 °58'W
Táchira
Cárdenas
S ofLa Grita; also called Pmo. (del) Guama! or Pmo. El Guama!
Guaramacal, Pmo. (de, del)
3200
09°l2'N
70° 13'W
Trujillo
Boconó
6 km SE ofBoconó, 6 km W ofGuaramacal; only a very small part at very top is subpáramo
Guaraque, Pmo. de
2873
08°l5'N
71 °44'W
Mérida
Rivas Dávila & Tovar
10 km S of Tovar, 10 km E ofBailadores, NE ofPmo. Los Carreros and connected with it, W ofPmo. de Jují and connected with it; also called Pmo. de San Francisco ( 1); not true páramo
Guaimaral, Pmo. Guama, Pmo. Guama!, Pmo. (del, El)
See Pmo. Guama
or50'N
72°07'W
Táchira
Libertad & Bolívar
Sierra La Maravilla, E of San Rafael
3600
08°56'N
70°30'W
Barinas/Trujillo
?/Boconó
Also spelled Pmo. Curiguay or Guirigay; Pico Guirigay (3860 m) is in Trujillo
Gutiérrez, Pmo.
3615
08°05'N
71 °58'W
Táchira
Hato Viejo, Pmo. (de)
4200
08°52'N
70°46'W
Mérida
Miranda & Rangel
N ofPmo. Mucuchíes and connected with it
Hemández, Cerro Las
4200
08°46'N
70°59'W
Mérida
Rangel
W ofMucuchíes
Horna, Pico
3220
08°03'N
71 °35'W
Mérida/Táchira
Arzobispo Chacón & Rivas Dávila!Uribante
SW ofLibertad (Canaguá), part ofPmo. Guaimaral
Guarin, Pmo. Guiriguay, Pmo. (de)
S ofLa Grita, SE ofPmo. Guama! and connected with it
08°14'N
71 °29'W
Mérida
Arzobispo Chacón
Cord. de Mérida, E ofPmo. El Molino and connected with it
4942
08°34'N
70°59'W
Mérida/Barinas
Libertador/Pedraza
Sierra Nevada de Mérida, NE ofPico Bolívar
Jabón, Pmo. (del, de El)
3440
09°34'N
70°07'W
Trujillo/Lara
Carache/Morán
Cord. de Trujillo, NNW of Campo Elías, E ofLa Concepción, E and SE slopes ofPico El Turma!, E ofPmo. de (El) Turma! and connected with it; Pico Jabón (3530 m) is in Trujillo/Lara
Jirajara, Pmo.
2800
09°09'N
70°l5'W
Trujillo
Boconó
S ofBoconó
Huesos, Pmo. Los Humboldt, Pico Isidro, Pmo.
:» "" ~ 3:::: oCll
See Pmo. San Isidro
Jubiote, Pmo. de
See Pmo. Juviote
Judío, Pico El
See Pmo. de Pata de Judío
Judío, Pmo. (El) Juviote, Pmo. (de)
\0 \0
.:S
See Pmo. de Pata de Judío 2301
09°25'N
70°l9'W
Trujillo
Trujillo & Boconó
Cord. de Trujillo; also spelled Jubiote; not true páramo
::::; \0
~
N ame Laguna (de El) del Gallo Laguna Brava, Pmo. de La
Elev. (m) Latitude
Longitud e Division 1
Division 2
Notes
Mérida
Libertador
Between Pico Bolívar and Pico Espejo
3300
Mérida
Rangel
Sierra de Santo Domingo, part ofPmo. de Los Granates, between Rio Los Granates and Loma·de Paja
Libertador
Sierra Nevada de Mérida, páramo surrounding this lake
4070
Laguna Coromoto
3400
Mérida
Laguna El Suero, Pmo.
4190
Mérida
Laguna Grande, Pmo. de
3700
08°50'N
70°54'W
Mérida
Rangel
Sierra de Santo Domingo, near Pmo. de Mucubají and Pmo. de Apartaderos
Laguna Negra, Pmo.
3500
08°47'N
70°48'W
Mérida
Rangel
Sierra de Santo Domingo, Mucubají area, near Pmo. de Mucubají and Pmo. de Apartaderos
Laguna Verde, Pmo.
4100
08°34'N
71 ooo·w
Laguna de Anteojos, Pmo. La
4000
08°30'N
Lagunq de Los Patos, Pmo. La
4200
Laguna las Canoas
3470
Laguna, Pmo. La
2800
Laja, Pmo. de La (2) Lajón, Pmo. El (1)
o>"Ij 5j
Mérida
Libertador
08°47'N
70°48'W
Mérida
Sierra de Santo Domingo, S ofLaguna Negra
08°48'N
70°48'W
Mérida
Sierra Nevada
08°18'N
71 °34'W
Mérida
Sucre & Tovar
15 km ESE ofTovar, S ofPmo. Monte Frío and connected with it, N ofPmo. Molino and connected with it; not true páramo
72°18'W
Táchira
Capacho
N ot páramo ( cloud forest)
71 °57'W
Táchira
Jáuregui
Sierra La Cimarronera, part ofPmo. La Cimarronera, NE of San José de Bolívar; also called Pmo. La Colorada
See Pmo. de Acequias
Lajas, Pmo. de Las 3620
or55'N
i0°56'N
Lajón, Pmo. El (2)
66°44'W
Not páramo (fide G. Morillo, pers. comm.)
Miranda
See Pmo. (de) Canaguá
Latalito, Pmo. 09°07'N
70°42'W
Trujillo
Valera
S ofLa Puerta
León, Pico El
4743
08°32'N
71 °06'W
Mérida
Libertador
SW part ofSierra Nevada de Mérida, NW ofLos Nevados
Leones, Pmo. (de) Los
4400
08°42'N
71 °10'W
Mérida
Andrés Bello & Libertador
Cord. del Norte (or de La Culata), N ofMérida, W ofMurucubá, E of Pmo. La Pedregosa and connected with it, SE of Pmo. El Salado and connnected with it, SW of Pmo. La Culata and connected with it (incl. La Lagunita and La Cañada Grande) NE ofLas Piedras
Laureles, Pmo. de Los
Linares, Pmo.
Trujillo
Llano Corredor, Pmo.
Mérida
Rangel
Mérida
Miranda & Justo Briceño W ofTimotes; incl. Pico El Fraile
Llano Grande, Pmo.
4200
09°00'N
70°49'W
~
Sierra Nevada de Mérida, Parque Nacional Sierra Nevada, toward Pico Bolívar
71 °06'W
or4S'N
~
til
o
Libertador
See Pmo. de. Acequias 1750
o
Sierra Nevada de Mérida, around Laguna Verde at the foot of Pico Humboldt
Mérida
Laja, Pmo. de La(!)
00
¡;¡;; IZl
til
ztil
::E ~ o
~ o
to
:;; ~
n
>
l'
C)
> ~ til
z
SSW ofPmo. de la Cristalina
<o r
00
_¡,..
Elev. (m)
Latitude
Longitude
Division 1
Division2
Notes
Llano del Trigo, Pmo. de
4000
08°40'N
70°52'W
Mérida/Barinas
Rangel/Bolívar
Sierra de Santo Domingo, N ofPmo. Los Pozones, NE ofPmo. Santo Cristo; incl. lagunas El Montón and Las Torres
Loma Negra, Pmo.
3000
08°09'N
71 °49'W
Mérida!Táchira
Rivas Dávila/Uribante
Loma Redonda, Pmo. Estación
4045
08°33'N
71 a04'W
Mérida
Libertador
Loro, Pmo. (de, del, El) (1)
4500
08°40'N
70°55'W
Mérida
Loro, Pmo. El (2) ·
4000
08°35'N
70°5l'W
Barinas
Pedraza
Táchira
Ayacucho
Táchira
Jáuregui
Sierra del Bata11ón, SE ofLa Grita, between Las Porqueras and Los Mutas, rd. to Queniquea
Mérida!Truji11o
Miranda & Justo Briceño/Escuque
NW ofTimotes; also ca1led Pmo. Alto Paramito
N ame
Sierra Nevada de Mérida, Parque Nacional Sierra Nevada, station on the Mérida Teleférico
SSE ofPmo. Los Pozones and connected with it See Pmo. (de, del, El) Loro (1)
Machado, Pmo. 3878
Malpica, Pmo. Manatará, Pico
· 1O km S of Bailadores, 18 km NNW of Pregonero, S part of Pmo. Las Tapias, E ofPmo. de Viriguaca and connected with it
Sierra Nevada de Mérida, 22 km E ofMérida; also ca1led Pmo. de Los Loros, Pmo. Loros, and Pmo. del Oro
Loros, Pmo. (de Los) Malpaso, Pmo.
\0 \0
.':S
09°04'N
70°49'W
3630
Manga, Pmo. La Maravilla, Pmo. (La)
2855
Mariño, Pmo. (de)
3517
08°!9'N
Masparro, Pmo.
3372
09°03'N
Media Luna, Pmo. de (la)
4230
Zulia
Colombia/Venezuela border
Trujillo
N ofValera
Táchira
Torbes & Uribante
Sierra LaMaraviUa; not true páramo
71 a49'W
Mérida/Táchira
Tovar & Rivas Dávila
Sierra del Bata11ón, 6 km W ofTovar, 10 km N ofBailadores, N ofPmo. La Negra
70°2l'W
Trujillo/Barinas
Boconó/Obispos
Ramal de Calderas
Mérida
Libertador
Sierra Nevada de Mérida, Parque Nacional Sierra Nevada, near Pico Bolívar, near the Teleférico Estación Loma Redonda
Mérida, Pmo. (de)
> "'
S:
:::::
o
(/)
Sierra Nevada de Mérida; probably a general terrn for many páramos near Mérida
Mérida
Mesa de Los Pozos
See Pmo. Mesa del Caba11o
Mesa, Pmo. de la
Truji11o 08°47'N
71°03
Carache
From Mesa de Carache on rd. toward Pmo. del Jabón
Mesallo, Pmo.
4672
Mesitas, Pmo. de Las
3300
Trujillo
Mérida
Micarache, Pmo. de
3000+
Mérida
Sierra de Santo Domingo?
Mifafi, Pmo. de
4500
Mérida
Sierra de la Culata, upper Mifafi Va11ey, part ofthe larger Pmo. de Piedras Blancas, south part ofPmo. de Mucuchíes
Mifés, Pico
4630
Mérida
Sierra de Santo Domingo
Above Niquitao Colinda with Pmo. El Rosario
;;;
Name
Elev. (m)
Latitude
Longitude
Division 1
Division 2
Notes
;;;; N
Mijará, Pmo. de
3300
Mérida
Arzobispo Chacón
15 km NNE ofChacantá
Minanón, Pmo.
3665
Mérida
Arzobispo Chacón
Between San José andEl Cupís; may egua! Pmo. de San José; sometimes spelled Minamón
Minugú, Pmo. de
3400
Mérida
Rangel
lO km SE ofSan Rafael de Mucuchíes, Quebrada Las Escaleras watershed
Miranda, Pmo. (de)
3600
08°57'N
70°49'W
Mérida
Miranda & Justo Briceño W ofChachopo, N ofPmo. Mucuchíes; part ofthe larger Pmo. Piedras Blancas
Misinta, Pmo. (de)
4000
08°47'N
70°55'W
Mérida
Sierra del Norte (orCulata), NE ofLa Culata
09°20'N
70°20'W
Trujillo
E ofTrujillo (not shown on maps)
t:Il
o ¡;:; ¡z¡ o"rl
Misisí, Pmo. (de) Mitisús, Pmo.
3724
Mérida
Rangel
Ramal de Sierra de Los Granates
Mocáz, Pmo. de
3100
Mérida
Libertador
Sierra Nevada de Mérida, 15 km NE ofEI Morro, 3 km S of Pico El Léon
Mérida
Arzobispo Chacón
Also spelled Macomboco
Mocosóz, Pmo.
3647
Mérida
Molino, Pmo. (del) El(!)
3270
Arzobispo Chacón, Sucre, Tovar & Campo Elías
25 km ESE ofTovar, S ofChiguara, E ofPmo. Río Negro and connected with it by Cerro Picacho, S of Pmo. La Laguna and connected.\Vith it, W ofPmo. El Serrucho
Mocomboco, Pmo. 08°l5'N
71 °33'W
Mérida
3859
Táchira
Uribante
Sierra La Cimarronera
Monigote, Pmo.
3670
08°57'N
70°41'W
Mérida
Miranda
Cord. de Trujillo, E ofTimotes
Monsalves, Pmo. de Oos)
3900
08°42'N
71 °08'W
Mérida
Montaña, Pmo. La
2840
08°12'N
71 °35'W
Mérida
Rivas Dávila & Arzobispo Chacón
22 km SE ofTovar, 30 km NE ofPregonero; true páramo?
Mérida?
Montañas del Morro 2840
Monte Plano, Cerro
3688
Monte Redondo, Pmo. de
2650
08°l9'N
71 °34'W
Mérida
70°ll'W
Trujillo
Monte de Palmar
Mérida
Moral, Pmo. El
Trujillo
Moraleño, Pmo. de
1973
Tovar & Sucre
Zulia 09°30'N
07°47'N
72°09'W
Táchira
...,
::e:
zt:Il
< o--<
~
t:O
o..., >z ñ >r Cl
See Pmo. del Morro
Monte Frío, Pmo.
~
t:Il
Molino, Pmo. El (2) Mono, Pmo. del
~
10 km E of Santa Cruz, 18 km E of Tovar, S ofPmo. Las Porqueras and connected with it, N of Pmo. La Laguna and connected with it; not páramo
>-
§ t:Il z
Colombia/Y enezuela border, Sierra de Perijá; called Cerro del Avión in Colombia Carache
Cord. de Trujillo, E of Santa Ana, NW ofCampo Elías; also spelled Monteredondo; not true páramo N ofValera
San Cristóbal
Sierra La Maravilla, E of San Cristóbal; not true páramo
<o r
00 -~'>-
N ame
Elev. (m) Latitud e
Longitude Division 1
Morita, Pmo. (de) La
3220
Trujillo
Morro, Pmo. (del)
2500
Mérida
Morro, Pmo. El
4037
08°33'N
71 °!2'W
Mérida
Division2
Notes
Boconó & Urdaneta
21 km from Jajó, 5 km before Tuñarne Near Mérida; also called Montañas del Morro; perhaps egua! to Pmo. El Cocuy (B. Briceño, pers. comm.)
CampoElías & Libertador
Sierra Nevada de Mérida, S of La Punta, SE of Ejido; also called Alto El Morro or Alto del Pmo. del Morro; does this egua! Pmos. (del) Morro? N ofPmo. El Volcán and connected with it; incl. Pico Guirigay (highest elevation)
Sierra de Palo Grande
Mortiño, Pmo. Cerro
2801
Motumbo,Pmo.de
3869
08°58'N
70°32'W
Trujillo/Barinas/Mérida
Boconó/Bolívar/Rangel
Mucamabás, Pmo. (de)
3975
08°56'N
70°44'W
Mérida
Miranda & Justo Briceño Rd. out of Trujillo, W of Chachopo; al so called Pmo. Mucumbás or Pmo. Mucumubás
Mucubají, Pmo. de
3700
08°47'N
70°48'W
Mérida
Rangel
Mucuchache, Pmo. La
3000+
08°47'N
70°50'W
Mérida
Mucuchachí, Pmo. de
2900
08°40'N
71 °20'W
Mérida
Mucuchíes, Pmo. (de)
3940
08°52'N
70°49'W
Mérida
Táchira
Sierra de Santo Domingo, 13 km NE of Mucuchíes, 25 km SSW ofTimotes Sierra de Santo Domingo
Miranda & Rangel
Union ofCord. de La Culata and Sierra Nevada de Mérida, 20 km SW of Timotes, 30 km NE of Mérida; incl. Pmo. de Las Cruces, Pmo. de Chachapos, Pmo. de Hato Viejo & Alto de Timotes, and Pmo. (Picacho El) Gavilán
"O
>~ ~ o IZl
See Pmo. (de) Mucamabás
Mucumbás, Pmo. Mucumpiche, Pmo.
4420
Mucumpisito, Pmo.
4539
08°54'N
70°53'W
Mérida
Sierra de Santo Domingo
Mérida
Justo Briceño & Miranda NW ofPico El AguiJa; misspelled as Mucumposito
Mucumposito, Pmo.
See Pmo. Mucumpisito
Mucumubás, Pmo.
See Pmo. (de) Mucamabás See Pmo. de Mucuñugue
Mucuñugue, Pico Mucuñugue,Pmo.de
4672
Mucuguí, Pmo.
3000
Mucurubá,Pmo.de
4100
08°18'N
71 °24'W
Mérida!Barinas
Rangel/?
Sierra de Santo Domingo; also called Pico Mucuñugue (fide G. Morillo, pers. comm.)
Mérida
Arzobispo Chacón & Campo Elías
W ofPmo. Las Coloradas and connected with it Sierra del Norte, NE ofMérida; not páramo (fide G. Morillo, pers. comm.)
Mérida
See Pmo. Mucutisís
Mucutusí, Pmo. Mucutisís, Pmo.
"' "'
~
4653
08°53'N
70°54'W
Mérida
Rangel & Justo Briceño
Sierra de La Culata, N of Apartaderos, NW ofPico El AguiJa; also called Pmo. Mucutusí
:;; V-l
N ame
Elev. (m) Latitud e
Longitude
Division 1
Division2
Notes
Mucuy, Pmo.
4470
Muerto, Pmo. El
3000
08°21'N
71 o 12'W
Mérida
Arzobispo Chacón & Campo Elías
26 km SSW ofMérida
Muertos, Pmo. Los
3112
or5TN
72°06'W
Táchira
Grita & Uribante
Sierra de Callejón Colorado, N ofPmo. Callejón Colorado and connected with it, NW of Queniquea
Mují, Pmo. de
3000
08°16'N
71 °42'W
Mérida
Rivas Dávila & Tovar
10 km SSE of Tovar, W part of Pmo. Río Negro, E ofPmo. Guaraque and connected with it, W of Pmo. La Zorra and separated from it by Río Agua Caliente
Mérida
~
tri
Muruti, Pmo. de
~
Mérida 2765
!0°34'N
66°50'W
o
Miranda/D.F.
V areschi (1954-1955) refers to the very top as páramo, but I have not included it as such
Nariz, Pmo. La
2962
09°40'N
70°07'W
Trujillo/Lara
Carache/Morán
Al so called Portachuelo de La Nariz
Negra, Pmo. (de) La
3400
08°15'N
71 °52'W
Mérida/Táchira
Rivas Dávila/Jáuregui
Sierra del Batallón, near Bailadores, between Tovar and La Grita, 30 km ofPregonero; also called Alto Pmo. de La Negra; sorne say 3080 mor 3658 m
Negro, Pmo.
!0°27'N
69°27'W
Lara
See Pmo. (de) Los Ñepes
Ñepes Alto, Cerro
See Pmo. (de) Los Ñepes
Ñepes, Pmo. (de) Los
3130
09°44'N
70°06'W
Lara/Trujillo
Morán/Carache
S ofSerranía Curarigua-Barbacoas; not páramo although maps say páramo; incL Pmo. del Ñepe Pequeño and Cerro Alto Ñepes
Nevados, Pmo. Los(!)
3500
08°30'N
71 °03'W
Mérida
Libertador
Sierra Nevada de Mérida, S slopes of Pico Espejo and Pico Bolívar; also called Sierra del Gallo
08°45'N
69°50'W
Mérida
Nevados, Pmo. de Los (2) 2500
Mérida
Niquitao, Pmo. de
SW part ofSierra Nevada de Mérida, connected with Pmo. Don Pedro, 25 km ENE ofMérida Sucre
Trujillo
48 km S ofEstánques, N ofPmo. (de) Canaguá
3240
08°l4'N
71 °57'W
Táchira
Panamericano & Jáuregui
1Okm N ofLa Grita, NE ofPmo. Los Pinos and connected with it, S ofPmo. de SanTelmo and connected with it
Novillo, Pmo. El
3320
08°55'N
71 °05'W
Mérida
Justo Briceño & Andrés Bello
30 km NE ofMérida, N ofPmo. Tucaní and connected with it
Oca, Pmo. de La
3500
09°34'N
70°06'W
Trujillo/Lara
Carache/Morán
Ongaravecoc, Pmo. de
Mérida 2400
Táchira
[/l
..,o >-3
;:¡: tri
ztri
:;E
o ~ to o
;:z
o>l'
Cl
>-
§ ztri
See Pmo. de La Teta de Niquitao
Nirgua, Pmo. de
Ocucuquí, Pmo.
:;!
~
Iribarren
Ñepe Pequeño, Pmo. del
Nieves, Pmo. Las
.¡o.
Sierra de Santo Domingo; 401 Om?
Munucuchimá, Pmo. de Naiguatá, Pico de
00
SW ofHumocaro Alto, connected with Pmo. Jabón Near Pueblo Nuevo
<o ¡---' 00
.¡o.
N ame
Elev. (m)
Latitud e
Longitude
Division 1
Division2
Oro, Pmo. (del) El (1)
Notes See Pmo. (de, del, El) Loro(!) See Pmo. (de, del, El) Loro (1)
Oro, Pmo. del (2) Ortiz, Pmo.
3481
09°03'N
70°24'W
Trujillo/Barinas
Boconó/Obispos
Ramal de Calderas, S ofNiquitao, N ofCalderas, NE ofPmo. de Calderas and connected with it, SW ofPmo. Castillejo and connected with it
Oso, Pico El
4566
08°46'N
71 °20'W
Mérida
Libertador & Andres Bello
Cord. de La Culata (or del Norte), W of Mucuchíes, N of Tabay; also called Alto del Oso
Oso, Pmo. El
2200
or47'N
72°08'W
Táchira
Cárdenas & San Cristóbal
Sierra La Maravilla, connected with Pmo. Moraleño and Pmo. El Pino; not true páramo
Oso, Pmo. del
2944
or52'N
71 °56'W
Táchira
Jáuregui & Sucre
Osorio, Pmo. de
3658
Táchira
Otlata, Pmo.
5 km SE of San José de Bolívar; does not equal Pmo. El Oso Along Tovar to La Grita rd.
Mérida
Pabellón, Pmo.
2300
or58'N
71 °33'W
Mérida!Táchira
Arzobispo Chacón/Uribante
Paiva, Pmo. de
2000
08°2l'N
71 °40'W
Mérida
Tovar
Pajarito, Pmo. El Pajarito, Pmo. de Los
SE ofMesa de Quintero, S ofGuaimaral (incl. Pico Pabellón), S ofPmo. de Altamira SW of Santa Cruz; not true páramo See Pmo. de Los Pajarito
3450
Palmas, Pmo. Las Palmira, Pmo. de
3300
Pan de Azúcar, Pmo.(!)
4710
Pan de Azúcar, Pmo. (2)
4000
Paramillo, Cerro El
09°04'N
70°33'W
Trujillo
09°42'N
70°08'W
Trujillo
Carache
Mérida
Miranda
Cerro Caracacho, between La Palma and Las González
Mérida
Justo Briceño & Libertador
Sierra del Norte, headwaters ofRío Mucujún, below Pico Pan de Azúcar
Boconó & Urdaneta
18 km SW ofNiquitao, S ofPmo. del Tostado and connected with it; also called Pmo. El Pajarito
08°50'N
71 o03'W
2600
08°13'N
71 °12'W
Mérida
Libertador
N ofCampo Elías; true páramo?
Paramito
1600
08°01'N
72°13'W
Táchira
Ayacucho
Village; not páramo
Paramito, Alto El
3640
09°03'N
70°5l'W
Mérida
Justo Briceño & Miranda 15 km W ofLa Mesa, 18 km S ofMonte Carmelo
Paramito, Cerro
2400
08°!0'N
71 °58'W
Táchira
Jáuregui
Paramito, Cerro El
1920
09°12'N
70°43'W
Trujillo
Es cuque
WSW ofEscuque; not true páramo
Paramito, El (1)
3200
09°02'N
70°48'W
Mérida!Trujillo
Miranda
Town along the Jajá-La Morita rd., toward Tuñame; not páramo
Paramito, El (2)
2800
Táchira
Junín
SE ofBetania, at the foot ofPmo. de Tamá; subpáramo at best
Paramito, El (3)
920
Zulia
Trujillo
"O
> ~ s;: oen
Near Michuntuí
5 km N ofLa Grita; not true páramo
Sierra de Perijá, San José de los Altos; not true páramo
Paramito, Pmo. El Páramo, El (1)
\0 \0
.:::¡
See Pmo. El Cocuy 2250
09°27'N
70°!9'W
Trujillo
Trujillo
S of Santa Ana, 15 km NE ofTrujillo; not true páramo 00 V>
N ame Páramo, El (2)
Elev. (m)
Latitud e
Longitud e Division 1
Division 2
Notes
1600
09°48'N
69°32'W
Lara
Jiménez
Locality SSE of Quibor, ENE of Gubiro; nota páramo Village W ofSanta Cruz; nota páramo
Páramo, El (3)
600
08°25'N
71 o42'W
Mérida
Bolívar
Páramo, El (4)
1600
or53'N
72°l4'W
Táchira
Lobatera
Páramo, Loma (del) El
1000+
10°!2'N
70°45'W
Zulia
Páramo, Montaña El
2200
09°38'N
69°44'W
Lara
Parros, Pmo. de Los
4400
Mérida
Pata de Judío, Pmo. de
3372
TáchiralApure
Pérez, Pmo. de
Village; not a páramo
Morán
NE ofGuarico, S ofEI Tocuyo, SSE ofEmba!se Dos Cerritos; not true páramo Sierra Nevada de Mérida, SE ofMérida, SW ofPmo. de los Loros
Cordoba!Páez
Colombia/Venezuela border, 30 km S (above) ofSan Vicente de la Revancha, E ofEI Cobre, SE ofPmo. de Tamá; same as Pmo. (El) Judío and Pico El Judío See Pmo. La Mucuchache
4240
2400
08°4l'N
08°04'N
71 o!3'W
71 °48'W
Mérida
Táchira
Campo Elías & Libertador
Sierra del Norte, N of Mérida, E of Pmo. Campanario and connected with it, S of Pmo. de Los Conejos and connected with it, SW of Pmo. El Salado and connected with it, W of Pmo. Los Leones imd connected with it
Uribante
5 km NW ofPregonero; not true páramo
Pico AguiJa, Pmo.
See'Pmo. i:lel.Aguila
Pico Bolívar, Pmo.
See Pico Bolívar
Pico La Garza
See Pico La Concha See Pmo. Piedras Blancas
Piedra Blanca, Pmo. Piedras Blancas, Pmo. (de)
4762
08°5l'N
70°56'W
Mérida
Justo Briceño & Miranda Sierra del Norte, 25 km NW ofTimotes, 10 km NNE of Mucuchíes, W ofPmo. Los Buitres and connected with it [incl. Pico Piedras Blancas (4778 m), Pico Los Nevados, Filo Los Nevados and Filo El Oso]; also called Pmo. Piedra Blanca; a biological reserve
Piedras, Pmo. de Las Piñango, Pmo. (de)
Probably equals Pmo. Piedras Blancas 4315
08°58'N
o,
Not true páramo
Pedregal, Pmo. de Pedregosa, Pmo. La
00
70°52'W
Mérida
Justo Briceño
Sierra de La Culata, S ofPiñango, N ofPmo. Las Tapias Sierra La Maravilla, N of Pico La Maravilla; not true páramo
Pino, Pmo. (d)el
2313
or4o'N
72°08'W
Táchira
Cárdenas
Pinos, Pmo. Los
2720
08°l2'N
71 °59'W
Táchira
Panamericano & Jáuregui 7 km N ofLa Grita, SW ofPmo. de Nirgua and connected with it
Pinos, Pmo. de Los
3000
08°25'N
71 °20'W
Mérida
Pintado, Cerro
3300
l0°30'N
72°50'W
Zulia
Piñuelas, Pmo. Las
3100
08°53'N
71 °23'W
Mérida
s;:: m s;::
o ;:a tZl ..,o ....,
::r: m
zm ::E o--< ~ b:l o ....,
;¡,.
2:1
n;¡,. r
Cl ;¡,.
Elm z
Sierra del Norte (near S end), N of Jají, 20 km W ofMérida Perijá
Sierra de Perijá, Serranía Valledupar (the northem part ofthe Sierra), the highest peak is Monte Plano (3688 m)
<o r
00
~
N ame Porqueras, Pmo. de Las
Elev. (m)
Latitude
Longitude
Division 1
Division 2
Notes
3070
08°10'N
71 °57'W
Táchira/Mérida
Jáuregui & Uribante/Rivas Dávila
NE ofLa Grita
Portachuelo, Pmo. (El) (1)
3070
08°22'N
71 °03'W
Mérida
Libertador & Campo Elías
Sierra Nevada de Mérida, SW ofPmo. El Toro and connected with it, between El Morro and Acequias
Portachuelo, Pmo. (de) El (2)
3070
08°09'N
71 °55'W
Táchira
Jáuregui
NW of Queniquea, connected with Pmo. Zumbador and Pmo. La Colorada; coordinates approximate
Portachuelo de La Nariz
See Pmo. La Nariz
Potrero Nuevo, Pmo.
3600
Mérida
Rangel
Pozo Negro, Pmo. ·(de, del)
3250
Mérida
Arzobispo Chacón & Campo Elías
Between San José and La Beguilla, close to Pmo. Canaguá and Pmo. El Cupís, part ofthe larger Pmo. (de) San José (2)
Pozones, Pmo. Los
4200
08°37'N
70°53'W
Barinas
Bolívar or Pedraza
Sierra de Santo Domingo, NE of Mérida, E ofPmo. Santo Cristo and connected with it, S of Pmo. Llano del Trigo and connected with it
Pozos, Pmo. Los(!)
3800
08°41'N
70°56'W
Mérida
Pozos, Pmo. Los (2)
3500
08°37'N
70°37'W
Mérida
Pozos, Pmo. Los (3)
3250
09°15'N
70°23'W
Trujillo
Trujillo & Boconó
Táchira
Uribante
Pregonero, Pmo. de
In the Loma Los Monsalves (B. Briceño, pers. comm.) Does this equal Pmo. Los Pozos(!)? 1Okm W ofBoconó, S ofPmo. El Atajo, N of Pmo. El Corazón
09°03'N
70°33'W
Trujillo
Urdaneta & Boconó
SE ofTuñame
3800
08°55'N
71 °44'W
Mérida
Miranda
Cord. de Trujillo, E ofChachopo, NW ofPueblo Llano
Trujillo
Urdaneta
3596
09°l0'N
70°45'W
Trujillo
Escuque & Valera
Púlpito, Pmo. (de El)
3912
08°05'N
72°l8'W
Táchira
Jáuregui
Pumar,Pmo.
2800
09°l1'N
70°13'W
Trujillo
Boconó
Quebradita, Pmo. de La
4000
Providencia, Pmo. de Pueblo Llano, Pmo. Pueblo Nuevo, Pmo. Puerta, Pmo. (de) La
\0 \0
~
"">--
~ ;:::: o (/J
W ofLa Puerta See Pmo. (de El) Púlpito
Púlpito, Pico
08°l5'N
Quemado, Pmo. El
71 °44'W
Sierra del Batallón; SE ofLa Grita, WNW ofPregonero; ·incl. Pico El Púlpito
Mérida
Sierra Nevada de Mérida, between Loma Redonda and La Aguada
Mérida
Not true páramo?
Quinorá, Pmo. (del)
See Pmo. (de) Quirorá
Quirorá, Pmo. 08°20'N
71 °27'W
Trujillo
Boconó
Mérida
Sucre & Campo Elías
Quirorá, Pmo. (de)
3260
Rafael Urdaneta, Cerro
3426
Zulia
Recostón, Pico
3890
Táchira
Uribante
Reinosa, Pmo. La
3720
Trujillo/Mérida
Urdaneta/Miranda
08°57'N
70°40'W
NE ofPmo. El Molino, 1Okm S ofPueblo Nuevo, SE ofEstanques Sierra de Perijá; páramo? Cord. de Trujillo, E ofTimotes
:;;; __,
N ame
Elev. (m) Latitud e
Longitude Division 1
Division2
Notes
Rejo, Pmo. (de) El
3400
08°24'N
71°19'W
Mérida
Campo Elías
15 km SE ofSan José
Revancha, Pmo. (La)
3450
07"32'N
72°23'W
Táchira
Junín
Part of Pmo. de Tamá area, SE of Delicias, Pico Revancha is 3457 m; also called Pmo. Rovancha
Táchira
San Cristóbal
Notpáramo
Rincón, Pmo. El 2032
Río Negro, Pmo. (de, del)
3223
08°l7'N
71 °39'W
Mérida
Rivas Dávila & Tovar
10 km SE ofTovar, S ofSanta Cruz, W ofPmo. Molino and connected with it by Cerro Picacho, E ofPmo. Guaraque and connected with it; incl. Pmo. de Mují and Pmo. La Zorra
Rivas, Pmo. de Los
3330
09° lO'N
70°44'W
Tmjillo
Escuque & Valera
SW ofMendoza, ESE ofMonte Carmelo, N ofPmo. San Rafael and connected with it
Rojas, Pmo. de
3160
09°00'N
70°25'W
Barinas
Bolívar
N of Las Calderas, S of Niquitao, S of Pmo. Calderas and connected with it, E ofPmo. Bartolo and connected with it
Rollal,Pmo.
See Pmo. El Royal 3890
08°02'N
0
7l 58'W
Táchira
Uribante & Jáuregui
1Okm N ofSan José de Bolívar, S ofLa Grita, W ofPregonero (incl. Pico ,El Rosal), S of Pmo. Batallón, W of Pmo. Cimarronera and connected with it, E of Pmo. La Tigra and connected. with it
Rosario, Pmo.
3560
08°ll'N
71 °55'W
Táchira!Mérida
Jáuregui/Rivas Dávila
8 km NE ofLa Grita, 12 km SW ofBailadores, N ofPmo. del Batallón and connected with it, SW of Pmo. La Negra and connected with it
Rosario, Pmo. (El, Los)
2700
09°l9'N
70°06'W
.Trujillo/Portuguesa
Boconó/Sucre
SSE ofCampo Elías, W ofBiscucuy, W ofSan Rafael de Palo Alzado, NE of Pmo. de Agua Fría; also <::alled Pmo. Los Rosarios; not true páramo; coordinates approximate
Rosas, Pmo. (de) Las
3245
09°35'N
70°07'W
Trujillo/Lara
Carache/Morán
Cord. de Trujillo, 11 km N ofLa Concepción, W ofHumocaro, slopes ofPico Cajingó, N ofPmo. de Turma! (1)
Rovancha, Pmo. Royal, Pmo. El
00
See Pmo. de Timotes
Río Frio, Pmo.
Rosal, Pmo. (de) El
;;;
See Pmo. (La) Revancha 4170
Sabana, Pmo. de La
1784
Sal, Pmo. de La
3910
08°40'N
70°50'W
Táchira
Capacho
09°0l'N
70°49'W
Mérida
Miranda & Justo Bríceño E ofPiñango
Mérida
Sierra de Santo Domingo-Sierra Nevada de Mérída union; also called Pmo. Rolla!
S:: tJ:j S:: o ¡;a [j]
o
'rj
....;¡
::r: tJ:j
z
tJ:j
::E
o-< ~ ttl o....;¡
>-z
ñ
>-
t-< Cl
>-
§ tJ:j z
Notpáramo
Salado Negro, Pmo.
1800
07"48'N
72°2l'W
Táchira
Libertad & Bolívar
Salado, Pmo. (El)
4486
08°42'N
71 °ll'W
Mérida
Libertador, Campo Elías Cord. del Norte (or de La Culata), N ofMérida, S ofPmo. de & Andrés Bello Los Conejos, W ofPmo. Los Leones and connected with it, E ofPmo. La Pedregosa
SW oflndependencia; not true páramo
<o r
..,.
00
N ame Salvaje, Pmo. (del, El)
Elev. (m) 3795
San Antonio, Pico de
Latitude
Longitud e Division 1
Division 2
09°06'N
70°35'W
Trujillo
Urdaneta
E of Jajó; may be referred to as Cerro Gordo, Pico Pajarito, or Fila El Paramito
Boconó
N ofPmo. Guaramacal, E ofBoconó, S ofPmo. Agua Fría
Sucre
E ofLa Trampa; not true páramo?
09°14'N
70° IO'W
Trujillo
San Antonio, Pmo.
3658
08°40'N
Mérida
San Bailón, Pmo.
2400
08°33'N
71 °03'W 71 o25'W
Mérida
San Francisco, Pmo. (1)
'D 'D
~
See Pmo. de Guaraque
San Francisco, Pmo. (2)
3445
San Isidro, Pmo.
2873
Táchira 08°2l'N
71 °33'W
Mérida
Tovar & Sucre
San José de Acequias, Pmo. de
NW of Pmo. El Molino, SW ofEstanques; also called Pmo. Isidro; not páramo See Pmo. (de) San José (2)
San José del Páramo
See Pmo. San José (1)
San José, Pmo. (1)
1450
San José, Pmo. (de) (2)
3250
San Pedro, Pmo. San Rafael, Pmo.
Notes
3600
Táchira
Capacho
Above La Laja; also called San José del Páramo; not true páramo
08°20'N
71 °l6'W
Mérida
Campo Elías, Libertador E of San José, above Veguilla (La Beguilla); incl. Monte El & Arzobispo Chacón Cupís and Pmo. del Cupís; also called Pmo. de San José de Acequias; may equal Pmo. Minanón
08°ll'N
71 °46'W
Mérida
Rivas Dávila
SE ofBailadores; part ofPmo. La Tapias?
09°08'N
70°45'W
Trujillo
Escuque & Valera
SW of Val era, W of La Puerta, SE ofMonte Carmelo, N of Pmo. Torres and connected with it, S of Pmo. Los Rivas and connected with it
Jáuregui& Panamericano
"O
>-
~ s;: o (IJ
Sierra del Batallón, 15 km N ofLa Grita, W ofPueblo Hondo, on Pico San Telmo (3265 m), N of Pmo. de Nirgua and connected with it; also called Pmo. La China; true páramo?
SanTelmo, Pmo. (de)
3259
08°16'N
71 °57'W
Táchira
Santa Bárbara, Pmo.(!)
3000?
or27'N
72°20'W
Táchira
Santa Bárbara, Pmo. de (2)
4252
Mérida
Santa Cruz, Pmo.
1400
Táchira
Santa Rita, Pmo. de
2200
Trujillo
Santo Cristo, Pmo. (de, del)
4200
08°37'N
70°57'W
Mérida/Barinas
Libertador & Rangel/Pedraza
Sierra Nevada de Mérida, E ofMérida, 20 km S ofMucuchíes; incl. Laguna Santo Cristo
Santo Domingo, Pmo.(!)
3373
or34'N
72°05'W
Táchira
Junín & San Cristóbal
Pmo. de Tamá area, NE ofPico El Cobre; also called Pico Santo Domingo
Santo Domingo, Pmo. (2)
4672
08°46'N
70°47'W
Mérida
Rangel
Between Río Santo Domingo and Río Chama, S of Pmo. Mucuchíes; incl. Pico Sto. Domingo or Mucuñuque at 4609 m
Seco, Pmo.
3290
Lara
Between Buenos Aires and Pmo. de Las Rosas
Secreta, Pmo. de La
2900
Táchira
NearFátima
NNW ofPmo. de Judío Between Pmo. de Tuñame and Pmo. de Timotes (a long distan ce) Capacho
Above Tres Esquinas; not true páramo Above Santa Rita; not true páramo
;;;; 'D
N ame
Elev. (m) Latitude
Longitude
Division 1
Division2
Notes
71 °29'W
Mérida
Sucre & Arzobispo Chacón
E ofPmo. El Molino; true páramo?
08°02'N
71 °59'W
Táchira
Jáuregui & Cárdenas ?
Divisoria Grita-Uribante, SE ofEl Cobre
09°05'N
70°47'W
Trujillo
Escuque & Valera
SW de La Puerta, S ofMonte Carmelo (incl. Pico El Perol, Pico El Rematadero, Pico San Pedro, Pico Escalera, Laguna La Gata and S s1opes ofCerro Las Cruces), N ofPmo. Llano Grande, S ofPmo. de Torres
7! 0 31'W
Mérida
Arzobispo Chacón
Táchira
Uribante
Serruchos, Pmo. (El, Los)
3245
08° 16'N
Sierrita, Pmo. La
3325
Siete Lagunas, Pmo. de (Las)
3894
Sinfonía, Pmo. La
2920
Ü8°11'N
Sombrero, Pmo.
2945
Somosica, Pmo.
14 km NE ofMesa de Quintero, bounded by Río El Molino on W and Quebrada El Barro on E; true páramo?
3860
08°02'N
71 °56'W
Táchira
Jáuregui & Uribante
12 km SSE ofLa Grita, E ofPmo. El Rosal and connected with it, N of Pmo. La Cimarronera and connected with it; also spelled Somosica
2850
Táchira
Jáuregui
Tabor, Pmo.
2847
Táchira
Junín
Nc;ar Colombia border; true páramo?
Tamá, Pmo. de
3500
or25'N
72°26'W
Táchira
JunínJApure
ColombiaNenezuela border
Tambor, Pmo. (de, del, El)
3054
08°36'N
71 °24'W
Mérida
Campo Elías, Sucre & Andrés Bello
Sierra de La Culata, NW of Jají, S ofLa Azulita, SW ofMérida
Tapias, Pmo. Las(!)
3517
08°12'N
71 o48'W
Mérida/Táchira
Rivas Dávila/Uribante
SE ofBailadores; incl. Pico Tapias at 3532 m
Tapias, Pmo. Las (2)
4200
08°58'N
70°53'W
Mérida
Justo Briceño
SW ofChachopo, W ofTimotes, N ofPmo. Mucumpisito
Tenegá, Pmo.
2972
or57'N
71 °56'W
Táchira
Jáuregui & Uribante?
Sierra La Cimarronera, E ofSan José de Bolívar
Tenerife, Pmo.
3202
Tenoca, Pmo.
Mérida
True páramo? ESE ofTrujillo
09°07'N
70°26'W
Trujillo
Urdaneta & Boconó & Ribas
Tetari, Pmo. del
3400
10°03'N
72°58'W
Zulia
Perijá
Tetas, Pmo. de las
4000
Tigra, Pmo. La
3600
08°0l'N
72°00'W
Tetari, Macizo de
o
tiJ
o
'Tj
--3
::e:
tTl
ztTl ~
Trujillo 4005
¡:;: tTl ¡:;: ~
See Pmo. Sumusica
Tablones, Pmo. Los
Teta de Niquitao, Pmo. de La
o
See Pico El Toro
Silla de El Toro
Sumusica, Pmo.
\0
Cord. de Trujillo, 29 km SSW of Trujillo; lower part is called Pmo. Carimbú
>-<;
o ~ to o--3
;¡,.
z
ñ ;¡,. t"'
a;¡,.
EltTl z
See Pmo. del Tetari
Trujillo Táchira
Sierra de Perijá, Serranía de Los Motilones (southem part of the Sierra), W of Machiques, along border with Colombia; incl. Pico Tetari at 3750 m; sometimes spelled Tetaria; also called Macizo de Tetarí SSW ofPmo. de la Cristalina
Jáuregui & Sucre
14 km S ofLa Grita, W ofPmo. El Rosal and connected with it
<o r
00
~
N ame
Elev. (m)
Latitud e
Longitud e
Division 1
Division 2
Notes
Timotes, Pmo. de
4286
08°52'N
70°50'W
Mérida
Miranda
Incl. Pmo. El Rincón (B. Briceño, pers. comm.) and Pmo. de Chachapos; also called Alto (de) Timotes; see also Pmo. de Mucuchíes
Tomón, Pmo. de
2400
09°13'N
70°43'W
Trujillo
Escuque & V aJera
Cord. de Trujillo, W ofMendoza; also called Alto de Tomón; not true páramo
Tonojó, Pmo.
2880
Trujillo
Trujillo & Boconó
EofTrujillo, alongTrujillo-Boconórd.; sometimes spelled Tonoso See Pmo. Tonojó
Tonoso, Pmo.
See Pmo. El Toro
Toro, Pico El Toro, Pmo. El
3600
08°25'N
71 °06'W
Mérida!Barinas
Libertador/Pedraza
Sierra Nevada de Mérida, S ofMérida, SW ofPico Bolívar; sometimes called Silla de El Toro or Pico El Toro?
Torotejopa, Cerro
3200
Zulia
Sierra de Perijá
Torrecita, Pmo. La
4547
Mérida
NNE of Sierra Nevada de Mérida toward Santo Domingo
Torres, Pmo. de (Los)
3680
09°07'N
70°45'W
Trujillo
Es cuque & V al era
W of La Puerta, N of Pmo. Las Siete Lagunas and connected with it, S ofPmo. San Rafael and connected with it
Tostado, Pmo. del
3800
09°06'N
70°33'W
Trujillo
Urdaneta & Boconó
20 km S ofSantiago de Trujillo, 12 km E ofJajó, N ofPmb. El Pajarito and connected with it, W ofPmo. Chorro Blanco and connected with it
Trampa, Pmo. La
2100
Tres Esquinas, Pmo.
2000
Mérida 08°2l'N
71 o52'W
Táchira
Not true páramo Panamericano
14 km WNW ofTovar; not true páramo See Pmo. El Trigo
Trigal, Pmo. El Trigo, Pmo. El
4170
09°00'N
70°49'W
Mérida
Miranda & Justo Briceño Sierra de La Culata, W ofTimotes; sometimes called Pmo. El Trigal?
Tucaní, Pmo.
4718
08°33'N
71 °40'W
Mérida
Andrés Bello, Libertador Cord. del Norte (or de La Culata), NNE ofMérida, NW ofPmo. & Justo Briceño Agua Blanca and connected with it, NE ofPmo. La Culata and connected with it, S ofPmo. El Novillo and connected with it; sometimes spelled Tucami ?
Tulata, Pmo. de la
'O
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(/)
Type of Stelis atra (Linden 687)
Mérida
Tunama, Pmo. de Tuñame, Pmo. (de)
\0 \0
::S
See Pmo. de Tuñame 3860
09°05'N
70°32'W
Trujillo
Urdaneta & Boconó
Cord. de Trujillo, W ofLas Mesitas, SE ofLa Quebrada; does this equal Pmo. El Salvaje?
Tuno, Pmo. del
3350
08°2l'N
71 °22'W
Mérida
Campo Elías
SW ofMérida, N ofPmo. Las Coloradas and connected with it
Turma!, Pmo. de
3600
09°30'N
70°08'W
Trujillo
Carache & Boconó
W and SW slopes, NW of Campo Elías, SE ofCarache, W of Pmo. Jabón and connected with it, E ofPmo. Monte Redondo and connected with it, SW ofPmo. Cendé; incl. Pico El Turma!
Turrnero, Pmo. El
4550
08°56'N
70°50'W
Mérida
Miranda & Justo Briceño Sierra de La Culata, W ofChachopo
::0
Elev. (m)
Latitude
Longitude
Tusta, Pmo. de
2200
08°24'N
71 °33'W
Urbina, Pmo.
1760
N ame
Division 1
Division2
Mérida
Sucre
Trujillo
S ofChiguara, NE ofTovar, E ofPmo. Las Porqueras
Mpio. Trujillo
Uricagua, Pmo. de
2700
Mérida
Vagabunda, Pmo. de la
2850
Mérida
Libertador
Along the Filo de la Vagabunda, El Morro-Aricagua rd., near El Portachuelo
Vela, Pico
3000
Táchira
Junín
Next to Pmo. de Tamá
Vichú,Pmo.
09°22'N
70°24'W
Trujillo
Vieja, Pmo. La (1)
08°5l'N
70°59'W
Mérida
Rangel & Justo Briceño
Cord. del Norte (or de La Culata), S ofTorondoy
09°07'N
70°28'W
Trujillo
Urdaneta & Boconó
E side ofTeta de Niquitao
Vieja, Pmo. La (2)
4000
Vieja, Pmo. La (3)
2773
~
Viejo, Pmo.
1200
or55'N
72°21'W
Táchira
Libertad
Viriguaca, Pmo. (de)
3421
08°10'N
71°5l'W
Táchira!Mérida
Uribante/Rivas Dávila
70°33'W
Mérida!Barinas
ti:I
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o
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Táchira?
08°56'N
\0 N
Ramal del Uribante; not true páramo
Táchira
Urbina, Pmo. La
Notes
Rangei/Bolívar
Village; not páramo 10 km S ofBailadores, 15 km ENE ofLa Grita, N ofPmo. del Batallón, W ofPmo. Las Tapias and connected with it, W of ·Pmo. Loma Negra and connected with it Origin of!Uo Aracay, N ofSanto Domingo
Volcán, Pmo. (El)
3700
Zerpa, Pmo.
4411
Zorra, Pmo. La
3000
08°15'N
71 °39'W
Mérida
Rivas Dávila
E ofBailadores, 15 km SE ofTovar, E ofPmo. de Mují; part of Pmo. Rio Negro?
Zumbador, Pmo. (del, El)
3250
or59'N
72°04'W
Táchira
Jáuregui
45 km SW of Tovar, N of Pmo. Almorzadero and connected with it, S of El Cobre; also called Pmo. Almorzadero (1) or Pmo. de Los Colorados
Sierra de LÓs:Granates
Mérida
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1999]
PARAMOS
193
Ecuador James L. Luteyn and Mauricio R. Gavilanes A.
The "Sierra" or montane part of Ecuador has two major parallel ranges of the Andes mountains, the Cordillera Occidental and Cordillera Oriental, between which is a relatively high interandean plateau averaging 2650 m. Severa! transverse mountain chains known as "nudos" connect the cordilleras and divide the intermountain plateau into roughly 1O basins or "hoyas." The Nudo del Azuay, at 4500 m, divides the country into two subregions, a northem area of modero volcanism andan area ofmore ancient volcanism to the south. The mountains in the north are higher, wetter, and their vegetation (including the páramos) generally resembles that of Colombia; whereas the mountains ofthe south are lower, drier, and their vegetation resembles that of northem Peru. The Sierra has at least 22 peaks averaging over 4200 m and 30 peaks of volcanic origin (six still active), giving rise to the region's moniker, "Avenue of the Volcanos." Estimates of the area covered by páramo in Ecuador vary considerably, depending on how páramo is defined, and range from 4470 km2 (Cañadas Cruz, 1983) to 19,61 Okm2 (Bonifaz, 1981; using subpáramo between 3500 m and 4000 m= 13,310 km2 and páramo above 4000 m= 6,300 km2) to 20,000 km2 (Encalada, 1986) toa high of25,000-28,000 km 2 (Acosta-Solís, 1984), out of a totalland area of ca. 280,000 km2 • [The figure of "about 40,000 km 2" given in Balslev and de Vries, 1991, was a mistake and should read "about 20,000 km2" (Balslev, pers. comm.).]
Ecuadorean national parks that include páramo are Cotopaxi, Podocarpus, and Sangay. Ecological reserves that include páramo are: Cotacachi-Cayapas, CayambeCoca, El Angel, and Antisana (cf. Mena V., 1995). The Ecuadorean gazetteer was difficult to assemble, because non e of the maps and índices consulted used the word "páramo" (in sorne cases the word "loma" seemed to be the equivalent, but not always). Therefore, the gazetterwas constructed in two ways: 1) from herbarium labels, local knowledge, and literature references that specifically used the word "páramo"; and 2) by the inclusion of al! localities found in geographical dictionaries, índices, and so forth that were over 3500 m and used the word "cerro," "laguna," "nevado," or "vulcán." That opened the possibility that sorne localities might not actual!y be páramo, but that was the decision made for the purposes of this book. And finally, as was mentioned in the introduction to the gazetteer, no secondary political divisions were used for the Ecuador gazetteer, and therefore the column "Division 2" was omitted from this table. For addítional discussion about Ecuadorean páramo vegetation, see Acosta-Solís, 1984; Balslev & de Vries, 1982, 1991; Black, 1982; Cerón M., 1985, 1992; Diels, 1937; J0rgensen & Ulloa U., 1994; Keating, 1995; León Yánez, 1993; L0jtnant & Molau, 1982; Mena V., 1984; Muñoz et al., 1985; and Ramsay, 1992.
Elev. (m)
Latitude
Longitude
Division 1
Notes
Achasipungu, Cerro
4120
01 °34'S
78°51'W
Bolívar
Cord. Occidental, 9 km NW of San Juan
Achayandi, Cerro
4000
00°57'S
78°55'W
Cotopaxi
Cord. Occidental, 8 km SE of Pi Jaló
Achilig, Cerro
4090
01 °35'S
78°49'W
Bolívar
Cord. Occidental, 7 km NW ofSan Juan
N ame
Achipungu, Cerro
4630
02° 14'S
78°32'W
Chimborazo
Cord. Oriental, 15 km NE ofTotoras
Achupallas, Cerro
3840
01 °11'S
78°43'W
Tungurahua
Cord. Oriental, 6 km NW ofQuisapincha
Achupallas, Pmo. de
3420
00°06'S
78°20'W
Pichincha
Cord. Oriental, also called Quebrada AguiJa
Aera Cruz, Cerro
3980
00°55'S
78°27'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 7 km NW ofLaguna de Los Anteojos
Aera, Cerro
3440
02°32'S
78°47'W
Cañar
1Okm NE oflngapirca
Ahuarongo, Cerro
4200
02° 19'S
78°45'W
Chimborazo
Cord. Oriental, 6 km of Achupallas
Ajos Pamba, Cerro
3880
01 o 49'S
78°51'W
Chimborazo
18 km SW ofSicalpa
Alalanda Chico, Cerro
4000
01 °55'S
78°3l'W
Chimborazo
Cord. Oriental, 15 km SE ofCebadas
Alalanda Grande, Cerro
4220
01 °55'S
78°30'W
Chimborazo
Cord. Oriental, 16 km SE of Cebadas
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Alambre Pamba, Cerro
3960
02°26'S
78°53'W
Cañar
9 km NE of Juncal
Alao, Pmos. de
4000
01 °49'S
78°35'W
Chimborazo
Cord. Oriental, W slopes V?lcán Sangay, near Cebadas, along rd. from Pungalá to Huancabamba
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Allcuquiro, Pmos. de
3622
02°39'S
78°30'W
Cañar
Cord. Oriental (Cord. de Allcuquiro), NE of Azogues; also spelled Allcuquiru; coordinates approximate
o;:>;!
Alpachaca, Pmo. de
3760
03 °30'S
79° 15'W
Azuay
Cord. Occidental, SE of Chaucha; also spelled Allpachaca
Alpacorral, Cerro
3600
78°59'W
Chimborazo
Cord. Occidental, 19 km NW ofVolcán Chimborazo
Altar Mayor, Cerro
3400
01 °25'S 03 o 32'S
o...;¡
79°22'W
Loja
4 km W of Selva Alegre
Altar, Cerro (El)
5319
01 o 41'S
78°24'W
Chimborazo/Morona-Santiago
Cord. Oriental, highest peak is Capa-urcu (incl. Laguna Pintada)
ñ > r
Altares, Pmo. de Los
4000
Chimborazo
Collanes Valley
Cl
Amaluza, Pmo.
3420
04 o 43'S
79°26'W
Loja
Cord. de Las Lagunitas, W of Jimbura
§
Amanta Grande, Cerro
3650
00°44'S
78°57'W
Cotopaxi
Cord. Occidental, 8.6 km SW of Sigchos
z
Amina Chiquito, Cerro
3995
00°57'S
78°23'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 2 km NE ofLaguna de Los Anteojos; also called Amina Chico
Amina Grande, Cerro
4188
00°58'S
78°23'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 1.5 km E ofLaguna de Los Anteojos
Amopuñuna, Cerro
4200
01 °2l'S
78°46'W
Chimborazo
Cord. Occidental, 11 km NE ofVolcán Chimborazo
Amorgeo, Pmo. de
3300
03 °09'S
78°5l'W
Azuay
Cord. Oriental; Sampedro (1 976) places it at 03 o !6'N/78 o 55'W on the Azuay/ Morona-Santiago border
Amoshayana, Cerro
4040
01 o 35'S
78°5l'W
Bolívar
Cord. Occidental, 15 km NE ofSan Simón
Añaguango, Cerro
4000
00°53'S
78°29'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 13 km NW ofLaguna de Los Anteojos
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N ame
Elev. (m)
Latitud e
Longitude Division 1
Notes
Angahuana, Cerro
3800
01 °20'S
78°42'W
Tungurahua
Cord. Occidental, 5 km SW ofTisaleo
Angamarca, Pmos. de
4000
78°50'W
Cotopaxi
Cord. Occidental, NE of Angamarca
Angasamana, Cerro
3400
01 °06'S 01 o23'S
78°30'W
Cotopaxi
Cord. Oriental, 9 km NE of Salcedo
Angel, Pmos. del
4000
00°43'N
7r54'W
Carchi
Nudo de Los Pastos, El Angel-Tulcán rd.; also called Pmo. de El Angel
Angochagua, Pmo. de
3800
78°01'W
Animullo, Cerro
4320
00°16'N 01 o 23'S
78°55'W
Chimborazo
Cord. Occidental, 12 km NW ofVolcán Chimborazo
Ansa Urcu, Cerro
4160
02° 19'S
78°44'W
Chimborazo/Cañar
Cord. Oriental, 12 km SW ofTotoras, 20.5 km NW ofHuangra
Ansahuaico, Cerro
4200
02°25'S
78°49'W
Cañar
17 km NE of Juncal
Antisana, Pmo. del
5704
00°32'S
78° 12'W
Napo/Pichincha
Cord. Oriental, SW slopes ofVolcán Antisana, 13 km S ofPapallacta
Angoas, Pmos. de
Cotopaxi!Napo
Cord. Oriental
lmbabura
Cord. Oriental, SE ofVolcán Imbabura, NE ofvillage of Angochagua
'D 'D
~
Antzaran, Cerro
3880
02°47'S
78°47'W
Cañar
19 km E of Juncal
Apagua, Pmo. de
4200
00°54'S
78°56'W
Cotopaxi
Cord. Occidental (Cord. de Apagua), between Zumbagua and Pilaló
Apicocha, Cerro
3480
02°50'S
79° IO'W
Azuay
Cord. Occidental, NW ofLaguna Llaviucu
Arcos, Pmo. de
3552
03 °34'S
79°27'W
Loja
Cerro de Arcos, 16 km W of Selva Alegre
Arenal del Chimborazo
4000
Chimborazo
Cord. Occidental, NE side ofVolcán Chimborazo
:>
Arrayán, Cerro
3400
01 °28'S
78°32'W
Tungurahua
Cord. Oriental, 9 k.rn S ofHuambalo
S::
Atacazo, Pmo. de
4570
00°2l'S
78°36'W
Pichincha
Cord. Occidental, Volcán Atacazo (4570 m), 9.7 km NW ofTambillo and 12 km from San Juan, S of Quito; also spelled Atacaso
Atán, Cerro
4160
02° 18'S
78°44'W
Chimborazo
Cord. Oriental, 10 km SW ofTotoras
Chimborazo
Cord. Occidental, 17 km NE ofPallatanga
Atapo Huañuna, Cerro
4200
01 °53'S
78°50'W
Atapulo, Cerro
3902
00°5l'S
78°44'W
Cotopaxi
Cord. Occidental, 8 km NW ofSaquisilí
Atillo, Pmo. de
3460
02° ll'S
78°30'W
Chimborazo
Cord. Oriental, near Lagunas de Atillo, 17 km NE ofTotoras; sometimes misspelled Atilio
Atio, Cerro
4200
01 o 32'S
78°53'W
Bolívar
Cord. Occidental, 13.5 km NE ofGuaranda
Atocha, Pmos. de
4000
01 °08'S
78°44'W
Cotopaxi
Cord. Occidental, 9 km SSW ofCusubamba
Atugrumi, Cerro
3800
00°52'S
78°56'W
Cotopaxi
Cord. Occidental, 9.5 km NE ofPilaló
Auchi, Cerro
3800
78°5l'W
Cañar
20 km NE of Juncal
Ayaguacha, Cerro
3920
02°2l'S 01 o 47'S
78°49'W
Chimborazo
12 km SW ofSicalpamba
Ayapunga, Cerro
4699
02°20'S
78°32'W
Chimborazo/Morona-Santiago/ Cañar
Cord. Oriental, E of Chunchi, SE of Achupalla; also spelled Ayapungo and Ayapungu
Ayapungo, Cerro
See Cerro Ayapunga
Ayapungu, Cerro
See Cerro Ayapunga
.,
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N ame
Elev. (m)
Latitud e
Longitude
01 °34'S
78°55'W
Azotado, Cerro El
4000
Azufra!, Pmo. del
3500
Division 1
Notes
Bolívar
Cord. Occidental, 7.6 km NE ofGuaranda
'-0 0\
Carchi
Azul Chico, Cerro
4000
02°l5'S
78°30'W
Chimborazo
Cord. Oriental, 18 km E ofTotoras
Bacha, Cerro
4120
01 °39'S
78°49'W
Chimborazo
Cord. Occidental, 6 km SW ofSan Juan
Bahuiña, Cerro
3647
02° 17'S
78°52'W
Chimborazo
Cord. Oriental, 5 km SE ofChunchi
Baizan Mularumi, Cerro
3800
01 °35'S
78°48'W
Chimborazo
Cord. Occidental, 4.7 km NW ofSan Juan
Balsas, Cerro
3951
03 °05'S
79° 17'W
Azuay
Cord. Occidental, 8.3 km NW ofSan Fernando
Banderilla, Laguna La
3450
04°l6'S
79°06'W
Loja
12.3 km SE ofVilcabamba
Basancel, Cerro
4350
01 °43'S
78°30'W
Chimborazo
Cord. Oriental, 9 km NE ofChambo
Berba Buenapata, Cerro
3800
01 °09'S
78°42'W
Tungurahua
Cord. Occidental, 10 km S ofCusubamba
Bishipotrero, Cerro
3880
02°04'S
78°40'W
Chimborazo
6.7 km E ofPalmira
Bodoqueras, Cerro
3440
02°42'S
78°35'W
Azuay
Cord. Oriental, 5 km E ofLas Palmas
Boladero, Pmo.
4100
Carchi
Around Laguna de Carr
Boliche, Altos de
3800
00°42'N
77°45'W
Carchi!Nariño (CO)
Ecuador/Colombia border, at foot ofVolcán Chiles; also called Nudo de Boliche, Pmo. de Boliche, or Nudo de Huaca (Cerro Boliche)
Boliche, Cerro
3840
01 °l2'S
78°54'W
Bolívar
Cord. Occidental, 10.3 km NE of Simiatug
Boliche, Nudo de
See Altos de Boliche
Boliche, Pmo. de
See Altos de Boliche
Boquerón, Pmo. de
Pichincha
Cord. Oriental, part ofthe Nevado del Cayambe area
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Bosay, Cerro
3580
02°06'S
78°5l'W
Chimborazo
Cord. Occidental, 7 km NW ofTixán
21n
Botijasacha, Cerro
4000
01 °26'S
78°58'W
Chimborazo
Cord. Occidental, 15 km NW ofVolcán Chimborazo
l'
Buerán, Pmo.
3818
02°35'S
78°55'W
Cañar
Cord. Oriental, 4.3 !c..m SE ofCañar, slopes ofCerro Buerán
o;p.
Buey Potrero, Cerro
3640
01 °09'S
78°27'W
Tungurahua
Cord. Oriental, 6.8 km SE of San José de Poaló
Builpacana, Cerro
3800
01 °59'S
78°43'W
Chimborazo
Cord. Oriental, 17 km E of Pallatanga
Elt"rl
Bulloc, Cerro
3760
01 °45'S
78°32'W
Chimborazo
Cord. Oriental, 8 km NE afLicto
02°05'S
78°50'W
Chimborazo
Cord. Occidental, 7.5 km NW ofTixán
Chimborazo
Cord. Oriental, E ofChunchi
Butoc Bailana, Cerro
3700
Cacheaco, Pmo. de
3800
Cagrin,Cerro
3836
01 °48'S
78°47'W
Chimborazo
12 km SW ofSicalpa
Cahuito, Cerro
4272
01 °07'S
78°43'W
Cotopaxi
Cord. Occidental, 7 km SSW ofCusubamba
Cailayacu, Cerro
3400
01 °5l'S
78°46'W
Chimborazo
4 km SE ofSan Jorge
Cajas, Cerro
4280
01 °38'S
78°52'W
Bolívar
Cord. Occidental, 11 km SW ofSan Juan
;p.
:z:
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N ame Cajas, Lagunas de
Elev. (m) 4020
Latitud e 00°39'S
Longitud e 78°2l'W
Division 1 Pichincha
Cord. Oriental, 14 km SE ofVolcán Sincholagua
~
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~
See Pmo. de Mojanda
Cajas, Nudo de Cajas, Pmo. (de, de Las, El Las, Las)
4100
Cajón de Quinuales, Cerro
4203
Calabozo, Cerro del
3760
Calacalí, Pmo. de
Notes
02°54'W
79°l7'S
Azuay
Cord. Occidental, ca. 30 km WNW of Cuenca and 3-10 km N of Soldados; possibly equals the Nudo del Azuay of Acosta-Solís (1984)
Ol 0 12'S
78°26'W
Tungurahua
Cord. Oriental, 8 km NE ofSucre
02°54'S
79°ll'W
Azuay
Cord. Occidental, 13 km W ofBaños
Pichincha
Doubtfully a true páramo (possibly a dry valley)
3680?
Calagüín, Cerro
See Cerro Calahuín
Calahuín, Cerro
3631
02°l2'S
78°54'W
Chimborazo
Calderón, Cerro
4240
01 °43'S
78°51'W
Chimborazo
10 km SE ofSicalpa
Calera Huaycu, Cerro
3600
01 o 40'S
78°55'W
Bolívar
Cord. Occidental, 8 km SE of San Lorenzo
Calera, Cerro La ( 1)
Cord. Occidental, W of Alausí; mispelled Calagüín
3800
01 o 43'S
78°32'W
Chimborazo
Cord. Oriental, 6.8 km E of Chambo
Calera, Cerro La (2)
3600
01 °5l'S
78°53'W
Chimborazo
Cord. Oriental, 16.6 km NE ofPallatanga
Callana Pucara, Cerro
4140
02° 17'S
78°47'W
Chimborazo
Cord. Oriental, 3.5 km from Achupallas
Calubín, Cerro
3463
02°10'S
78°58'W
Chimborazo
Cord. Occidental, 1O. 7 km NW of Sibambe
;¡,.
Cotopaxi
Cord. Occidental, 8.5 km NW of Saquisilí
S: ;;::
00°49'S
78°44'W
3530
03 °Ü5'S
78°45'W
Azuay
Cord. Oriental, 6.6 km SE of Sigsig
3271
00°13'N
78°27'W
Pichincha
Cord. Occidental, NW ofSan José de Minas
Campamento, Cerro El
3548
00° 12'S
78°32'W
Pichincha
Cord. Occidental, 7.5 km NE ofChillogallo, E ofQuito
Campana Pungu, Cerro
3600
Ü2°ll'S
78°56'W
Chimborazo
Cord. Occidental, 6.5 km NW ofSibambe
Campana Urcu, Cerro (1)
3400
00°37'S
78°47'W
Pichincha
Cord. Occidental, 7 km E of Santa Rosa
Campana Urcu, Cerro (2)
4360
01°l0'S
78°52'W
Cotopaxi
Cord. Occidental, 15.5 km NE ofSimiatug
Campana Urcu, Cerro (3)
4160
01 °04'S
78°54'W
Cotopaxi
Cord. Occidental, 5.5 km NE of Angamarca
Campana, Cerro
4200
01 o 41'S
78°50'W
Chimborazo
7.5 km NW ofSicalpa
02°57'S
79°20'W
Azuay
Cord. Occidental; may equal Quebrada Can Can (3600 m)
Cotopaxi
Cord. Occidental, NE slopes Volcán Illiniza, NW ofPastocalle
Calvario,. Cerro
3540
Calvario, Cerro El Cambugán, Pmos. de
Cangan, Pmo. de Caniyalo, Pmo. de
3500
Cantzag, Cerro
4023
01 °l0'S
78°40'W
Cotopaxi/Tungurahua
Cord. Occidental, 11 km SSE ofCusubamba, 7 km NE ofQuinzapincha, W ofPillaro
Capadia Chico, Cerro
4320
01 o 25'S
78°56'W
Chimborazo
Cord. Occidental, E ofVolcán Chimborazo
Capadia Grande, Cerro
4280
01 °25'S
78°56'W
Chimborazo
Cord. Occidental, 14 km NW ofVolcán Chimborazo
Caparina, Cerro ( 1)
3600
00°57'S
78°57'W
Cotopaxi
Cord. Occidental, 3.5 km SE ofPilaló
Caparina, Cerro (2)
3600
01 o 48'S
78°52'W
Chimborazo
Cord. Occidental, 17 km SW of Sicalpa
""'
o
en
~ _,
N ame
Elev. (m)
Latitud e
Longitude Division 1
Notes
Caparina, Cerro (3)
4000
02°05'S
78°38'W
Chimborazo
10 km SE ofPalmira
Caparina, Cerro (4)
3800
02°22'S
78°52'W
Cañar
15.5 km NE of Juncal
Caparosa, Cerro
3405
01 o 45'S
78°58'W
Chimborazo
Cord. Occidental, 8 km SE of Santiago
Carboncillo, Pmo. de
3000
03 °33'S
79°15'W
Azuay!Loja
Loma Carboncillo with highest point at 3067 m; not true páramo
Carboneria, Cerro
3800
01 °2l'S
78°43'W
Tungurahua
Cord. Occidental, 6.7 km SW ofTisaleo
Carga Allichina, Cerro
3547
02°35'S
78°50'W
Cañar
\0 00
12 km SE ofCañar See Pmo. Carihuairazo
Cariguayrazo, Pmo. Carihuairazo, Pmo. (de)
5028
01 °24'S
78°45'W
Tungurahua
Cord. Occidental, NE of Volcán Chimborazo; also called Cerro and spelled Cariguayrazo (Spanish) or sometimes Caryhuayrazo
S:: m S::
Carnero Pungu, Cerro
4194
78°46'W
Tungurahua
Cord: Occidental, 22 km NE of Simiatug
Carniceria, Cerro
4320
01 °!0'S 01 o38'S
78°52'W
Bolívar
Cord. Occidental, 10 km SW ofSan-Juan
~ en
Carniceria, Pmo. de
3750
Chimborazo
W ofVolcán Sangay
o>r:l
Carshau, Cerro
3800
Cañar
8 km NE of Juncal
::r: m
02°25'S
78°55'W
o
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zm
Casaguala, El Cerro
See Cerro Casalahua
Casahuala, Cerro
see Cerro Casahihua
:;;::
Tungurahua
Cord. Oriental, NW of Amqato; sometimes misspelled Casahuala, El Casaguala or Cazaguala
o ~ tJ:j o >! z ñ
Casalahua, Cerro
4545
01 oIO'S
78°45'W
~
Cascajo, Cerro
3832
02°58'S
79°22'W
Azuay
Cord. Occidental, 9 km SE of Chaucha
Cascarillas, Lagunas de
4000
79°!6'W
Azuay
Cord. Occidental, 15.6 km NE ofChaucha
Cashisho, Cerro
3760
02°52'S 01 ooo·s
78°29'W
Cotopaxi
Cord. Oriental, 10.4 km NNW of San José de Poaló
Caspiasana, Cerro
3455 4120
03 °09'S
79°02'W
Azuay
Cord. Occidental, 8.7 km SW ofCumbe
02°22'S
78°45'W
Cañar
5 km NE of Laguna Las Tres Cruces
Cl
Cord. Occidental, 9.4 km NW ofSan Juan
§ m z
Caspifitina, Cerro Caspigasí, Pmo. de
Pichincha 01 °33'S
Caspitotora, Cerro
4080
Castillo, Pmo. del
3400
Caucay, Cerro
3800
02°32'S
Cayambe, Pmo. de
3700
Cazaguala, Cerro
4100
78°50'W
Bolívar Azuay
Cord. Oriental, 6-8 km NNE of Sevilla de Oro
78°59'W
Cañar
6.7 km NW ofCañar
00°02'N
77°59'W
Pichincha/Imbabura/Sucumbíos Cord. Oriental, slopes ofVolcán or Nevado Cayambe (5789 m)
01 °09'S
78°45'W
Cotopaxi
Cazagua1a, Pmo. de
>-l' >--
Cord. Occidental, 19.3 km SE of Angamarca See Cerro Casalahua
Cazco Valenzuela, Pmo. de la comuna
3350
00°14'S
78°09'W
Imbabura
Cebadas, Cerro
3800
02°26'S
78°56'W
Cañar
5.2 km NE of Juncal
<o r
00
.¡,.
N ame Cenerón, Cerro
Elev. (m)
Latitude
4120
Chacana, Cerro
Longitude
Division 1
Notes
01 o 44'S
78°3I'W
Chimborazo
Cord. Oriental, 8 km E of Chambo
00°26'S
78°14'W
Pichincha
Cord. Oriental, 13 km SW ofPapallacta
Chagrapamba, Cerro
4000
01 °49'S
78°51'W
Chimborazo
17 km SW of Sicalpa
Cha1ata, Cerro
4200
01 °30'S
78°50'W
Chimborazo
Cord. Occidental, 14.5 km NW ofSan Juan
Cha1úa, Pmos. de
3400
00°48'S
78°44'W
Cotopaxi
Cord. Occidental, 6 km W ofCanchagua Grande; páramo?
Chambi, Cerro
4240
01 °06'S
78°46'W
Cotopaxi
Cord. Occidental, 17 km NE of Angamarca
Chame, Cerro
3600
00°56'S
78°52'W
Cotopaxi
Cord. Occidental, 12.6 km E ofPilaló
Chanchán, Cerro
3850
01 °49'S
78°47'W
Chimborazo
14 km S of Sicalpa
Chaning, Pmo. de
3500
Azuay
E of Azogues; a Lehmann collecting site; may equal Pmo. de Guairacaja Cord. Occidental, W ofGuamote, E ofPallatanga; also spelled Chanloor
\0 \0 \0
~
Chanlor, Cerro
4367
01 °52'S
78°50'W
Chimborazo
Chaparumi, Cerro
4000
01 o 44'S
78°30'W
Chimborazo
Cord. Oriental, 9.6 km SE ofChambo
Charonhuasi, Cerro
3840
03 °07'S
79°!9'W
Azuay
Cord. Occidental, 8.5 km NW of San Fernando
Charopungu, Cerro
4200
02 °22'S
78°50'W
Chimborazo
10 km SE ofCompud
Chaso Carapungu, Cerro
4283
01 °09'S
78°49'W
Cotopaxi
Cord. Occidental, 12 km SE of Angamarca
Chasquí, Pmo. de
3658
Chauchavi, Cerro
4148
01 °27'S
78°56'W
Bolívar
Cord. Occidental, 4 km NW ofQuindihua Grande
> ~
Chaulor, Cerro
4367
01 °54'S
78°50'W
Chimborazo
Cord. Occidental, W ofGuamote
o
Chaupiata, Cerro
3640
01°17'S
78°49'W
Tungurahua
Cord. Occidental, 15.3 km E ofSimiatug
Chaupirrumi, Cerro
3955
00°53'S
78°18'W
Napo
Cord. Oriental (Cord. de Llanganates), 13 km NE ofLaguna de Los Anteojos; also spelled Chaupirumi
Chaupiurcu, Cerro (1)
4400
01 °05'S
78°48'W
Cotopaxi
Cord. Occidental, 14 km NE of Angamarca, 6 km SE ofHuairapungo
Chaupiurcu, Cerro (2)
4240
02°22'S
78°47'W
Cañar
23 km NE of Juncal
Chaupiyacu, Cerro
4245
02°21'S
78°44'W
Cañar
Cord. Oriental, 19 km NW ofHuangra
Chepe1, Pmo. de
2950
ElOro/Loja
Cord. Occidental (Cord. de Chillán), NE ofZaruma, between Payama and Tioloma
Chico Tunyal, Cerro
4200
Cañar
21 km NE ofJuncal
Chihuila-Sacha, Pmo.
3700
Cotopaxi/Tungurahua
Cord. Oriental, part ofPmos. de Llanganati
Chilca, Pmo. de
4352
01 °03'S
78°49'W
Cotopaxi
Cord. Occidental, 12 km NE of Angamarca (incL Cerro Taquirrumi)
Chiles, Pmo. de
4768
00°48'N
7r55'W
Carchi
Nudo de Los Pastos, Volcán Chiles, 9.3 km NW ofTufiño, 34-3 6 km W of Tulcán
Chilindalo, Cerro
3400
00° 13'S
78°33'W
Pichincha
Cord. Occidental, 4.2 km NE ofLloa
Chiltazón, Pmo.
3967
00°41'N
78°01'W
Carchi
Cord. Occidental
Chimborazo, Cerro (1)
3508
01 °37'S
78°31'W
Chimborazo
Cord. Oriental, 4 km NE of Quimiag
Pichincha (León ?)
02°24'S
78°47'W
"O
~
(/)
::0 \0
N
Elev. (m)
Latitude
Longitude
Division 1
Notes
Chimborazo, Cerro (2)
3900
02°35'S
78°42'W
Cañar
Cord. Oriental, 13 km NW ofHuangra
Chimborazo, Pmo.
6310
01 °27'S
78°48'W
Chimborazo
Cord. Occidental, NW ofRiobamba; also called Volcán Chimborazo and Nevado Chimborazo (at 6310 m)
Chimobrazo, Loma
4014
00°28'N
78°20'W
Imbabura
Cord. Occidental, N ofNevado Cotacachi
Chinibano, Cerro
4180
00°56'S
78°28'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 8 km NW ofLaguna de Los Anteojos
N ame
Chique, Laguna
See Laguna Puruhanta
Chiquelín
May egua! Pmo. Llangagua
o o
See Laguna Puruhanta
;:;::
Chiqnicagna, Pmo. de
3800
01 °24'S
78°45'W
Tnngurahua
Cord. Occidental, NE slopes ofCarihuairazo, SW ofPilahuín; coordinates approximate
;:;::
Chiqningu, Cerro
3400
0! 0 49'S
78°45'W
Chimborazo
13.6 km S ofSicalpa
Chiquirignal, Cerro
4320
00°ll'S
78°34'W
Pichincha
Cord. Occidental, 7 km N ofLioa
Chiquita, Laguna
3800
00°07'S
78°!6'W
Pichincha
8 km NE ofTocachi
Chiqniurcn, Cerro
4000
01 °!3'S
78°48'W
Cotopaxi
Cord. Occidental, 18 km NE ofSimiatug
Chiripungu, Cerro
3440
02°25'S
78°56'W
Cañar
7.5 km NE of Juncal
Chiquí, Laguna de
t:Il
o
~
(/)
o'Tl
..., ::r: t:Il
zt:Il
:E '"< o
Chisag, Cerro
4320
01 °0l'S
78°52'W
Cotopaxi
Chitamachi, Cerro
4000
01 °36'S
78°5l'W
Bolívar
Cord. Occidental, 8 km NW of San Juan
Chivo, Cerro El
4080
01 °04'S
78°56'W
Cotopaxi
Cord. Occidental, 3 km NW ofAngamarca
~
Chocarcocha Chico, Laguna
4200
02° 15'S
78°3l'W
Morona-Santiago
Cord. Oriental, 14-16.5 km SE ofTotoras
o..., ;¡...
Tnngurahua
Cord. Oriental, 4 km NE ofSan José de Poaló
ñ ;¡...
Chorrera de Panza, Cerro
4080
Chozones, Pmo. de
3000
Laja
Between Oña and Saraguro; probably not true páramo
Chugchilán, Pmo. de
3500
Cotopaxi
Cord. Occidental, NW ofToacazo, NE slopes ofVolcán Illiniza
Pichincha
Coordinates estimated
Chnmillos, Pmo. de
3600
0
ÜÜ ]2'S
78°26'W
78°13'W
Chuquibanza, Cerro
May equal Pmo. Llangagua 02°32'S
78°59'W
Cañar
5.7 km NW ofCañar
4200
01 °!3'S
78°47'W
Tnngurahua
Cord. occidental, 19.3 km NE ofSimiatug
3800
00°37'S
78°32'W
Cotopaxi
Cord. Oriental, 1Okm SE of Chanpi
Chnrupucará, Cerro
3600
01 °l0'S
78°56'W
Cotopaxi
Cord. Occidental, 12.7 km NE ofSimiatug
Chusalongo, Lagunas de
3900
02°55'S
79°!5'W
Azuay
NW ofSoldados
Chnspilli, Cerro
4000
0! 0 47'S
78°53'W
Chimborazo
17.3 km SW of Sicalpa
Chusurrumi, Cerro
4000
01 °49'S
78°50'W
Chimborazo
16 km SW ofSicalpa
Chuqnirahua, Cerro (1)
3800
Chuquirahna, Cerro (2) Churupinto, Pmo. de
tD
See Laguna Chocarcocha Chico
Chocarcocha Grande, Laguna 01 °05'S
Cord. Occidental, 12 km NE of Angamarca; latitude may be 00°45'S instead (?)
z
t-
Cl
;¡, ?O t:l t:Il
z
<o r ""' 00
N ame
Elev. (m) Latitud e
Longitud e Division 1
Notes
4189
02°06'S
78°37'W
Chimborazo
11.7 km SE ofPa1mira
Cipreses, Laguna de Los
4200
02°50'S
79°!7'W
Azuay
Cord. Occidental, 14.3 km NE ofChaucha
Cisarán, Cerro
4360
02° !9'S
78°48'W
Chimborazo
Cord. Oriental, 6. 7 km SW of Achupallas
Clavelillo, Cerro
3880
02°55'S
79°!2'W
Azuay
. Cintura Pi terina, Cerro
'O
'D
~
Cord. Occidental, 23.5 km SW ofCuenca See Pmo. de Corredores
Clavelillo, Pmo. Cochaloma, Cerro
4063
01 °58'S
78°48'W
Chimborazo
Cord. Occidental, 17 km NE ofPallatanga
Cojitambo, Cerro
4120
02°25'S
78°47'W
Cañar
21.7 km NE of Juncal
Colacunde, Cerro
3600
00°45'S
78°47'W
Cotopaxi
Cord. Occidental, 12 km SE ofSigchos
Cola!, Cerro
4686
02° 15'S
78°32'W
Chimborazo/Morona-Santiago
Cord. Oriental, SE ofPalmira, E of Achupallas
Colay, Cerro
4685
02°!4'S
78°30'W
Chimborazo
Cord. Oriental, E of Alausí
Collapungu, Cerro
4240
or24's
78°48'W
Cañar
20.3 km NE of Juncal
Collincora, Cerro
3560
04 o28'S
79°19'W
Loja
15.2 km SE ofSan Antonio de Aradas
Común, Cerro
4400
01 °03'S
78°49'W
Cotopaxi
Cord. Occidental, 13.5 km NE of Angamarca
Conde, Cerro
4215
00°46'S
78°2l'W
Napo
Cord. Oriental, 13.5 km SE ofVolcán Cotopaxi
Cóndor Amana, Cerro
4302
78°45'W
Cotopaxi
Cord. Occidental, 19 km SE of Angamarca
Cóndor Machay, Cerro
4080
01 °08'S 01 o31'S
78°49'W
Bolívar
Cord. Occidental, 13 km NW of San Juan
Cóndor Matzay, Cerro
4200
01 °07'S
78°45'W
Cotopaxi
Cord. Occidental, 19 km SE of Angamarca
Cóndor Samana, Cerro
3600
00°58'S
78°58'W
Cotopaxi
Cord. Occidental, 4.7 km SE ofPilaló
Cóndor, Cerro
3850
Chimborazo
6.7 km SE ofPalmira
3988
02°04'S 01 o20'S
78°40'W
Condorcaca, Cerro
78°44'W
Tungurahua
Cord. Occidental, 7.3 km SW ofTisaleo
Condorrnatze, Cerro
4000
00°58'S
78°54'W
Cotopaxi
Cord. Occidental, 11 km SE ofPilaló; does this equal Cerro Cóndor Matzay?
Condorpalte, Cerro
4000
01 °34'S
78°50'W
Bolívar
Cord. Occidental, 9 km NW of San Juan
Condorpasana, Cerro
4240
01 °24'S
78°56'W
Bolívar
Cord. Occidental, 16 km NW ofVolcán Chimborazo
Condorpaso, Cerro
4197
00°57'S
78°25'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 4 km NE ofLaguna de Los Anteojos
Condoructo, Cerro
4199
01 °39'S
78°52'W
Bolívar
Cord. Occidental, 11 km SW of San Juan
Conga, Cerro
4434
00°52'S
78°16'W
Napo
Cord. Oriental, 2 km S ofLaguna Yanacocha (not to be confused with the Yanacocha on the N side ofVolcán Pichincha) Cord. Occidental, 20 km NW of San Fernando
Contrayerba, Cerro
4040
03°0l'S
79°23'W
Azuay
Copaloma, Cerro
3425
00°57'S
78°44'W
Cotopaxi
Cord. Occidental, 5 km W ofPujilí
Coparahuashicuna, Cerro
4200
78°49'W
Cotopaxi
Cord. Occidental, 12.5 km NE of Angamarca
Corazón Urcu, Cerro
3690
01 °03'S 01 o31'S
78°45'W
Chimborazo
Cord. Occidental, 12.6 km NE of San Juan
"O
> ~ s;: o (/]
N
o
N
N ame
Elev. (m)
Latitude
Longitud e
Division 1
Notes
Corazón, Cerro
4320
00°3l'S
78°39'W
Pichincha
Cord. Occidental, 11 km SW ofMachachi
Cordaurcu, Cerro
4160
01 o 45'S
78°53'W
Chimborazo
15.2 km SW ofSicalpa
Loja/Zamora-Chinchipe
Loja/Zamora-Chinchipe border, E ofSaraguro
Cordoncillo, Pmos. de Corral Pamba, Cerro
3800
00°43'S
78°49'W
Cotopaxi
Cord. Occidental, 7 km SE ofSigchos
Corral Viejo, Cerro
3720
ooo 19'S
78°35'W
Pichincha
Cord. Occidental, 6 km SW ofChillogallo
o
N
Corredores, Pmo. (de)
3260
03 o 33'S
79°35'W
El Oro
Cord. Occidental (Cord. de Chillán), N ofZaruma; also called Pmo. Clavelillo
Cotacachi, Pmo. de
3400
00°22'N
78°20'W
Imbabura
Cord. Occidental, 15 km E ofPlaza Gutiérrez (incl. Laguna Cuicocha and Nevado Cotacachi at 4937 m); also called Volcán or Cerro Cotacachi
Coto Amina, Cerro
4155
00°5l'S
78o28'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 15 km NW ofLaguna de Los Anteojos
Coto Juan, Cerro
4160
01 o 43'S
78°52'W
Chimborazo
11 km SW of Sicalpa
Cotopaxi, Volcán
5897
00°40'S
78°26'W
Cotopaxi/Pichincha/N apo
Cord. Oriental, 18 km NE ofMulaló, N ofLatacunga; also called Nevado Cotopaxi
Cresta de Gallo, Cerro
3520
04° 15'S
78o06'W
Loja
11.6 km ofVilcabamba
Criadero de Buitres, Cerro
3920
02°50'S
79°1l'W
Azuay
21 km NW of Cuenca
Cruz de Gula, Cerro
3690
03 °42'S
79°19'W
Loja
7.3 km NW ofSan Lucas
ztn
01 °0l'S
78°50'W
Cotopaxi
01 °17'S
78°26'W
Tungurahua
Cord. Oriental, 6.4 km SE ofSucre
o>-<;:o
Cruz, Pmo. de La Cruzloma, Cerro (1)
3770
o % o'Tl (/l
-3
::r: tn
~
Cruzloma, Cerro (2)
3450
01 °20'S
78°4l'W
Tungurahua
Cord. Occidental, 3.5 km W ~fTisaleo
Cruzloma, Pmo. de
3050
00° 15'S
78°34'W
Pichincha
Cord. Occidental, slopes ofVolcán Pichincha, above Quito
Cruzmaqui, Cerro
3700
01 o 20'S
78°43'W
Tungurahua
Cord. Occidental, 6.5 km W ofTisaleo
Cruzpungu, Cerro
4281
02°07'S
78°39'W
Chimborazo
9.2 km SE ofPalmira
Cuarteles, Cerro
3560
02°25'S
78°57'W
Cañar
7.6 km NE of Juncal
Cubijín Urcu, Cerro
3440
01 °02'S
78°29'W
Cotopaxi
Cord. Oriental, 0.6 km NE ofCashaloma
Cubilán, Cerro (1)
3880
03 °04'S
79° 18'W
Azuay
Cord. Occidental, 10 km NW ofSan Fernando
Cubilán, Cerro (2)
3767
03°34'S
79°23'W
Loja
6.2 km SW ofSelva Alegre
Cubillán, Cerro
4300
00°39'S
78°l4'W
Napo
Cord. Oriental, between Volcán Cotopaxi and Cerro Antisana, SE ofCerro Laurel
Cubillín, Pmo. de
3400
Chimborazo
Cord. Oriental, but also see Volcán Quilimás
;>:; tD
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ztn
See Volcán Quilimás
Cubullín, Volcán Cuchiguasi, Cerro
~ tn ~
4113
00°52'S
78°27'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 12 km NW ofLaguna de Los Anteojos
Cuchihuasi, Cerro
4460
01 °04'S
78°50'W
Cotopaxi
Cord. Occidental, 11 km NE of Angamarca
Cuiquiloma, Cerro
4206
01 o 33'S
78°5l'W
Bolívar
Cord. Occidental, 12 km NW ofSan Juan
Culebrillas, Pmo.
3450
02°53'S
78°46'W
Azuay
Cord. Oriental, 23--40 km SE ofGualaceo
<o ¡:-' 00
.¡:.
N ame
Elev. (m)
Culebrillas, Pmo. de
3600
Cumbas, Pmos. de
3800
Cumbaya, Pmo.
3450
Latitude
Longitude
Division 1
Notes Cord. Oriental, in Parque Nacional Sangay
00° 19'N
78°22'W
Imbabura
-
\0 \0
~
Cord. Occidental, N ofLaguna de Cuicocha; may egua! Pmos. de Cungas
Pichincha Napo (?)
Cord. Oriental, E slopes, E of(San Miguel de) Salcedo
Cunambay, Cerro
4000
01 °39'S
78°48'W
Chimborazo
Cord. Occidental, 5.2 km SW of San Juan; may egua! Cerro Cunumbay
Cungas, Pmo. de
4000
00°20'N
78°2l'W
Carchi
Cord. Occidental, SSW ofNevado Cotacachi, E ofPmos. de Tablachupa, W of Pmo. de Morochos; may egua] Pmo. de Cumbas
Cunumbay, Cerro
4085
01 o 40'S
78°48'W
Chimborazo
5 km NW ofSicalpa; may egua! Cerro Cunambay
Cunupugro, Cerro
3840
00°54'S
78°28'W
Cotopaxi
Cumbijín, Pmo. de
Curiquinga, Pmo. de
Cord. Oriental (Cord. de Llanganates), 10.5 km NE ofLaguna de Los Anteojos See Pmo. de Curiquingue
Curiquingue Y anarrumi, Cerro
3800
00°5l'S
78°5l'W
Cotopaxi
Curiquingue, Cerro ( 1)
4405
01 °04'S
78°48'W
Cotopaxi
Cord. Occidental, 14.7 km NE of Angamarca
Curiquingue, Cerro (2)
4080
01 o 13'S
78°46'W
Tungurahua
Cord. Occidental, 6.4 km NW ofHuilaló
Cañar
5 km S ofCañar, slopes ofCerro Buerán; also spelled Curiquinga
Cord. Occidental, 5.5 km E ofLaguna Quilotoa
Curiquingue, Pmo. de
3800
Cushiro, Cerro
3562
04 o 18'S
79°07'W
Loja
12.4 km SE ofVilcabamba
Cushorrumi, Cerro
3400
02°06'S
78°5l'W
Chimborazo
Cord. Occidental, 8 km NW ofTixán
Cushquisurcuna, Cerro
4200
01 °40'S
78°52'W
Bolívar
Cord. Occidental, 11 km NW of Sicalpa
Cushuna, Cerro
3637
01 °34'S
78°56'W
Bolívar
Cord. Occidental, 6.6 km NE of Guaranda
Cusín, Cerro
3990
00° ll'N
78°07'W
Cusnopaccha, Pmo. de
3400
Cusumunllo, Cerro
4340
01 °24'S
Cutahua, Cerro
3400
Daldal, Pmo. de
Imbabura!Pichincha
Cord. Oriental, N ofCayambe, NW ofOlmedo
Chimborazo
Cord. Oriental, E of Alao; also spelled Cushnipaccha
77°54'W
Bolívar
Cord. Occidental, 11 km NW ofVolcán Chimborazo
01 °l3'S
78°57'W
Cotopaxi
Cord. Occidental, 6.2 km N of Simiatug
3600
01 o 48'S
78°32'W
Chimborazo
Cord. Oriental; altitude and coordinates are for village of Daldal
Danas, Cerro
3778
02°09'S
78°53'W
Chimborazo
Cord. Occidental, 9 km SW ofTixán
Danas, Cerro El
4200
01 o 48'S
78°55'W
Chimborazo
Cord. Occidental, 14 km SE ofSantiago
Danta Sirina
3995
00°52'S
78°24'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), N ofLaguna de Los Anteojos
Derrumbo, Cerro El
3720
01 °48'S
78°53'W
Chimborazo
18 km SW ofSicalpa
Diablo Peña, Cerro
3840
01 o 41'S
78°53'W
Bolívar
Cord. Occidental, 11 km NE of Santiago
Diablo Sirina, Cerro
4520
02°16'S
78°32'W
Morona-Santiago
Cord. Oriental, 15 km SE ofTotoras
Dolliurcu, Cerro
4160
01 o 12'S
78°48'W
Tungurahua
Cord. Occidental, 19 km NE of Simiatug
Dudaloma, Cerro
3400
02°34'S
78°45'W
Cañar
Cord. Oriental, 20 km SE of Cañar
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N
ow
N
Name
Elev. (m)
Latitud e
Longitude
Division 1
Notes
o
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Dudas, Cerro
4031
02°24'S
78°37'W
Cañar
Cord. Oriental, 7.7 km NW ofHuangra
Encantada, Laguna
3900
Cord. Oriental, 8.6 km S ofOyacachi
4100
78°03'W 78o30'W
Napo
Ensillada, Cerro
00°!5'S 01 o47'S
Chimborazo
Cord. Oriental, 9 km NE ofLicto
Era Urcu, Cerro
4473
01 °0l'S
78o50'W
Cotopaxi
Cord. Occidental, 13 km NE of Angamarca
Escalera Grande, Cerro
3520
02°30'S
78°46'W
Cañar
17 km E ofTambo
Esmoquero, Cerro
4040
00°55'S
78°23'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 5 km N ofLaguna de Los Anteojos
Esperanza, Cerro La
3800
01 °03'S
78°26'W
Tungurahua
Cord. Oriental, 5 km NE of San José de Poaló
Estribo Grande, Cerro
4406
78°48'W
Cotopaxi
Cord. Occidental, 14.5 km NE of Angamarca
Eva, Cerro
3400
01 °04'S ooo 14'S
78°39'W
Pichincha
Cord. Occidental, 8 km NW ofLioa
Falto, Cerro
4210
ooo 57'S
78°27'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 7 Km W ofLaguna de Los Anteojos
Fiachi, Laguna
3800
02°53'S
79°09'W
Azuay
Cord. Occidental, 9 km NW ofBaños
o"rj
Fierro Urcu, Cerro
3788
03°4l'S
79°20'W
Loja/EI Oro
Cord. Occidental, WSW of Saraguro, 9 km NE of Gua! el
Fierroloma, Cerro
4137
03°02'S
79°22'W
Azuay
Cord. Occidental, 18 km NW ofSan Fernando
5ti
Foyo Huasi, Cerro
4080
01 °07'S
78°44'W
Cotopaxi
Cord. Occidental, 8.5 km SW ofCusubamba
z
See Pmos. de Mojanda
><:
Fuya-Fuya, Cerro
2:: tJ:j 2::
8
:>:1
(/l
tJ:j tJ:j
~
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Gaballushi, Cerro
3480
02°26'S
78°57'W
Cañar
4 km N of Juncal
Galápagos, Pmo. de
3600
02°42'S
78°35'W
Azuay/Morona-Santiago
Cord. Oriental, E ofEl Pan
Galgalán, Cerro El
4686
02°09'S
78°24'W
Chimborazo/Morona-Santiago
Cord. Oriental, E ofPalmira, WNW ofMacas
Gallo Rumi, Cerro
4500
02°23'S
78°50'W
Cañar
17.7 kmNEofJuncal; [Prov. Chimborazo, Cord. Occidental (fide Acosta-Solís, 1984)]
Gallo Urcu, Cerro
4095
01 °08'S
78°43'W
Cotopaxi
Cord. Occidental, 9 km SW ofCusubamba
ñ
Gallocantana, Cerro
3840
00°l9'S
78°36'W
Pichincha
Cord. Occidental, 9 km SW ofChillogallo, SW ofQuito
1:"" Cl
Galo Cocha, Laguna
3800
01 °l9'S
78°57'W
Bolívar
Cord. Occidental, 3.5 km SW ofSimiatug
Ganquis, Cerro
4058
01 °35'S
78°5l'W
Bolívar
Cord. Occidental, 10 km NW ofSan Juan
Gapaguey, Cerro
3480
01 °34'S
78°46'W
Chimborazo
Cord. Occidental, 6 km NE ofSan Juan
Garzaguachana, Cerro
3800
01 °54'S
78°54'W
Bolívar
Cord. Occidental, 9.5 km NE ofSan Simón
Chimborazo
SE ofSicalpa
0
Gavilán Huachana, Cerro
3909
Ol 48'S
78°47'W
Gavilán Paccha, Cerro
4160
02°20'S
78°46'W
Chimborazo
19 km E ofCompud
Gigantones, Cerro
4040
03 °0l'S
79°24'W
Azuay
Cord. Occidental, 22.3 km NW ofSan Fernando
00°45'S 01 o43'S
78°45'W
Cotopaxi
Cord. Occidental, may equal Pmo. de Güintza (Nudo de Tiopullo)
4200
78°5l'W
Chimborazo
Gintza, Pmo. de Gorda Urcu, Cerro Gorda, Cerro
10-12 km SW ofSicalpa; also called Cerro Gorda See Cerro Gorda Urcu
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N ame
Elev. (m)
Latitud e
Longitud e Division 1 Chimborazo
Notes Cord. Oriental, SE ofCebadas, W ofVolcán Sangay
Gosoy, Pmos. de
4190
01 °57'S
78°34'W
Grande de Mojanda, Laguna
3719
00°08'N
78° 15'W
Pichincha!lmbabura
1Okm NE ofTocachi; also called Laguna de Mojanda or Laguna de Mojanda Cajas
Grande, Laguna
3660
02° ll'S
78°46'W
Chimborazo
Cord. Oriental, 7 km NE of Alausí
Grande, Pmo.(!)
4060
00°33'S
78°31'W
Pichincha
Cord. Oriental, Cerro Rumiñahui, NW slopes, SE ofMachachi
00°29'S
78°39'W
Pichincha
Cord. Occidental, N slopes ofCerro Corazón, SW of Aloag; may egua! Pmo. El Pongo
Chimborazo
Grande, Pmo. (2)
3800
Guabo, Pmo.
3400
Guagra Huma, Cerro
4000
01 °36'S 01 o IO'S
78°46'W 78°27'W
Tungurahua
Guagrahuma, Cerro
3550
03°43'S
79°19'W
Loja
Cord. Oriental, 10.2 km NE ofSucre 7 km NE of Gua! el; al so called Nudo de Guagrauma See Cerro Guagrahuma
Guagrauma, Nudo de Guagua Pichincha, Pmo. del
\0 \0 \0
~
4875
ooo IO'S
78°36'W
Pichincha
Cord. Occidental, part ofVolcán Pichincha, W ofQuito, 8 km NW ofL!oa, 11.4 km SW ofNono; also spelled Huahua Pichincha
Guairacaja, Pmo. de
3500
Azuay
May egua! Pmo. de Chaning
Gualalí, Loma
4380
Imbabura
Cord. Occidental, Volcán Cotacachi
Gualampuz, Pmo.
3865
Guamallo, Cerro
3640
02°32'S
78°49'W
Cañar
5 km E oflngapirca
Guamaní, Pmo. de
4200
00°20'S
78°33'W
Pichincha!Napo
Cord. Oriental, E ofPifo on rd. to Papallacta; also called Monte Guamaní, El Paso de Guamaní, and Pmo. de la Virgen
Guangotasín, Cerro
4000
00°57'S
78°29'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 10 km NW ofLaguna de Los Anteojos
Guanguloma, Cerro
3750
Cañar
Cord. Oriental, 5.7 km SW ofHuangra
3816
02°29'S 01 o 28'S
78°39'W
Guaslán, Cerro
78°35'W
Tungurahua
9.7 km SE ofMocha
Guasumpala, Cerro
4000
01 °45'S
78°3l'W
Chimborazo
Cord. Oriental, 9 km SE ofChambo
Guingopana Grande, Cerro
4055
00°45'S
78°49'W
Cotopaxi
. Chimborazo
Cord. Occidental, Volcán Chimborazo, E slopes "t!
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5 km E oflsinlivi See Tiopullo
Güintza, Pmo. Gulag, Cerro
38oo
01 °38'S
78°49'W
Chimborazo
Cord. Occidental, 4.4 km SW of San Juan
Gulag, Pmos. de
4206
01 o 33'S
78°5l'W
Bolívar
Cord. Occidental, ENE ofGuaranda, NW ofSan Juan
Habacota, Laguna
3800
03 °Ü5'S
79°19'W
Azuay
Cord. Occidental, 9 km NW of San Fernando
Hermoso, Cerro
4571
01 o 13'S
78°17'W
Tungurahua
Cord. Oriental (Cord. de Los Llanganates), ca. 12 km SE of Pmo. de Jaramillo; also called Cerro Llanganati (M. Gavilanes, pers. comm.)
Hignug, Cerro
3400
01 o 41'S
78°42'W
Chimborazo
4.3 km W ofYaruguíes
Huacaguañuna, Cerro
3880
01 o 46'S
78°49'W
Chimborazo
Cord. Occidental, 7 km NE of Juan de Velasco
Huachana Curiguingue, Cerro
4080
00°5l'S
78°26'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 13 km NW ofLaguna de Los Anteojos
See Altos de Boliche
Huaca, Nudo de
N
o
V>
Name
Elev. (m)
Latitud e
N
Longitude Division 1
Notes
Huacorrumi, Cerro
4022
00°52'S
78°28'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 12 km NW ofLaguna de Los Anteojos
Huagra Huanuna, Cerro
3800
01 °49'S
78°49'W
Chimborazo
Cord. Occidental, 6 km E of Juan de Velasco
Huagra Moyuna, Cerro
3490
02°34'S
78°45'W
Cañar
13 km SE oflngapirca
Huagra Urcu, Cerro
3600
01 °50'S
78°46'W
Chimborazo
2.2 km E of San Jorge
Huagracorral, Cerro
3800
78°4l'W
Tungurahua
6.6 km NW ofQuisapincha
Huahua Pari, Cerro
4320
Ül 10'S 01 o 13'S
78°53'W
Cotopaxi
Cord. Occidental, 10-15 km NE ofSimiatug
Huahua Urcu, Cerro
3400
01 °5l'S
78°54'W
Chimborazo
Cord. Occidental, 16.5 km NE ofPallatanga
Huahuayacu, Cerro
4153
01 °l4'S
78°48'W
Tungurahua
Cord. Occidental, 16 km NE ofSimiatug; may equal Cerro Sombrero Urcu
0
Huahua Pichincha, Cerro
o
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See Pmo. del Guagua Pichincha
Huaico, Pmo.
See Pmo. Huayco
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Huaicutambo, Cerro
3440
01 °ll'S
78°29'W
Tungurahua
Cord. Oriental, 4 km N ofBaquerizo Moreno
Huailhuacer, Cerro
3800
01 o 36'S
78°47'W
Chimborazo
Cord. Occidental, 2.6 km SW ofSan Juan
Huairacajana, Pmo. de
3000
Azuay
Near Pindilig; not true páramo
Huairaja, Cerro
4007
02°06'S
78°38'W
Chimborazo
9 km SE ofPalmira
Huairapamba, Cerro
3800
01 o 47'S
78o47'W
Chimborazo
1Okm SW of Sicalpa
Huairapata, Cerro
3800
01 °l0'S
78°44'W
Tungurahua
9.7 km NW ofQuisapincha
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Huairapungo, Cerros de
4272
01 °38'S
78°52'W
Bolívar/Chimborazo
Cord. Occidental, W of San Juan, NW of Juan de Velasco; also spelled Huairapungu; coordinates approximate
?'
Huaitayug, Cerro
4160
01 °39'S
78°53'W
Bolívar
Cord. Occidental, 10.5 km SE ofSan Simón
:;¡,.
Hualicanga, Cerro
3960
02°25'S
78°55'W
Cañar
8.3 km NE of Juncal
Chimborazo/Morona-Santiago
Cord. Oriental
ñ:;¡,.
Huamboya, Pmos. de Huangu, Cerro El
3700
02°30'S
78°38'W
Cañar
17.4 km NW ofHuarainac
Huangupud, Pmos. de
4200
01 °45'S
78°5l'W
Chimborazo
SW ofSicalpa
Huapageo, Cerro
3495
03 °Ü8'S
79°03'W
Azuay
8.7 km SW ofCumbe
Huapungoto, Cerro
3520
01 °4l'S
78°56'W
Bolívar
Cord. Occidental, 7 km NE of Santiago
Huaquer, Pmos. de
3604
00°35'N
7r53'W
Carchi
Cord. Occidental, W of San Gabriel (incl. Loma de Huaquer)
Huarcorrumi, Cerro ( 1)
4000
01 °20'S
78°44'W
Tungurahua
Cord. Occidental, 8 km W ofTisaleo
Huarcurrumi, Cerro (2)
3880
ÜÜ 50'S
78°22'W
Napo
Cord. Oriental (Cord. de Llanganates), 15 km NE ofLaguna de Los Anteojos
0
Huarcurrumiloma, Cerro
3880
Cotopaxi
Cord. Occidental, 5 km SW of Cusubamba
3845
01 °06'S 00ol4'N
78°43'W
Huarrni Imbabura, Cerro
78°12'W
Imbabura
Cord. Oriental, NNE ofLago de San Pablo
Huatouiña, Laguna
3660
03 o 40'S
79°2l'W
Loja
11 km NE of Gua! el
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Elev. (m)
Latitud e
Huaycu, Pmo.
4000
01 o55'S
78°49'W
Chimborazo
Cord. Oriental, 12 km E ofGuamote; also spelled Huico
Huaytayuc, Cerro
3960
01 o40'S
78°54'W
Bolívar
Cord. Occidental, 10.5 km NE ofSan Lorenzo
Huerajachupa, Cerro
3840
02°05'S
78°39'W
Chimborazo
7.6 km E ofPa1mira
Huiñahuarta, Cerro
3600
78°39'W
Azuay
Cord. Oriental, 4 km SE ofTaday
Huiñatus, Cerro
4000
02°39'S 01 o39'S
78°48'W
Chimborazo
Cord. Occidental, 5 km SW ofSan Juan
Huisla, Cerro
3762
01 °23'S
78°34'W
Tungurahua
4.7 km SE ofQuero
N ame
Longitude
Division 1
Notes
-
\0 \0
~
See Pmo. de Tiopullo
Huinsa, Pmo. Humanchi, Cerro
4080
or52'S
79°13'W
Azuay
Cord. Occidental, 25.6 km W ofCuenca, 16 km W ofSayausí
Iglesia, Cerro
4360
01 °06'S
78°48'W
Cotopaxi
Cord. Occidental, 14 km E of Angamarca
Iglesia-Rumí, Cerro
3969
01 °36'S
78°49'W
Chimborazo
Cord. Occidental, 5.5 km NW ofSan Juan
Ignillapungu, Cerro
3840
78°53'W
Cañar
8.6 km NE of Juncal
Igualata, Pmo. de
4000
02°28'S 01 o29'S
78°38'W
Tungurahua
Cord. Oriental, 7.2 km SE of Yanayacu (incl. Cerro Igualata at 4330 m)
Iguila Pungu, Cerro
3665
02°32'S
78°50'W
Iguán, Cerro
4000
Probably equals Cerro Igua1ata in Tungurahua
Igualata-Sanancajas, Pmo. de . Cañar
4 km NE oflngapirca
Carchi
Iguán, Laguna
3625
02° 12'S
78°32'W
Chimborazo
Cord. Oriental, 15.5 km NE ofTotoras
Iliniza Norte, Nevado
5600
00°38'S
78°43'W
Pichincha/Cotopaxi
Cord. Occidental, 30 km W ofVolcán Cotopaxi, 1Okm SW ofChaupi, N ofLatacunga
Iliniza Sur, Nevado
5800
00°39'S
78°42'W
Pichincha/Cotopaxi
Cord. Occidental, 30 km W ofVolcán Cotopaxi, 10.5 km SW ofChaupi, N ofLatacunga
Illahuas, Cerros
4230
01°10'S
78°45'W
Tungurahua
Cord. Occidental, 25 km NE ofSimiatug
Imanante, Cerro
4200
01 °50'S
78°31'W
Chimborazo
Cord. Oriental, 14 km NE of Cebadas
Imbabura, Cerro
4630
00°15'N
78°10'W
Imbabura
Cord. Oriental, E ofOtavalo, 7 km N ofSan Pablo
Inabuela, Laguna
4200
01 °21'S
78°44'W
Tungurahua
Cord. Occidental, 8.7 km SW ofTisaleo
Inca Cocha, Lagunas
4000
or51'S
79°16'W
Azuay
Cord. Occidental, 15.6 km NE ofChaucha
Inga Casa, Laguna
4000
02°5l'S
79° 16'W
Azuay
Cord. Occidental, 16.6 km NE ofChaucha
Ingañán, Cerro
3730
02°29'S
78°45'W
Cañar
23.5 km E of Juncal
Inganilla, Pmo. de
3400
78°55'W
Cañar
Between Biblián and Cañar
Ingapirca, Cerro
4200
02°37'S 01 o44'S
78°30'W
Chimborazo
Cord. Oriental, 10 km SE ofChambo
Irarrumi, Cerro
4000
02°05'S
78°37'W
Chimborazo
11 km SE ofPalmira
Ishcurrumi, Cerro
3600
01°43'S
78°49'W
Chimborazo
6 km SW of Sicalpa
Ishquirrumi, Cerro
3860
01 °06'S
78°43'W
Cotopaxi
Cord. Occidental, 6 km SW ofCusubamba
lshunshico, Cerro
3400
01 °22'S
78°40'W
Tungurahua
Cord. Occidental, 3 km S ofTisa1eo
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N
o
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Name.
N
Elev. (m)
Latitud e
Longitud e
Division 1
Notes
Isicsilli, Cerro
4000
00°54'S
78°24'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 6.5 km N ofLaguna de Los Anteojos
Jaboncillo, Cerro
4120
02° 17'S
78°43'W
Chimborazo
Cord. Oriental, 7.6 km SW ofTotoras
Jacsán, Laguna
4000
02° 17'S
78°35'W
Chimborazo
Cord. Oriental, 10.3 km SE ofTotoras
Jalopamba, Cerro
3600
01°47'S
78°5l'W
Chimborazo
14 km SW ofSica1pa
Jalopungu, Cerro
3460
02°34'S
78°45'W
Cañar
13 km SE oflngapirca
Jaramillo, Pmo. de
4100
01 °l2'S
78°26'W
Tungurahua
Cord. Oriental (Cord. de Los Llanganates), ca. 12 km NW ofCerro Hermoso
Jatun,Muca, Cerro
4082
00°47'S
78°47'W
Cotopaxi
1O km SE oflsinlivi; also spelled Jatún Muea
J atunsamana, Cerro
3533
02°33'S
78°45'W
Cañar
14 km SE oflngapirca
Jichaisita, Cerro
4030
00°58'S
78°30'W
Cotopaxi
Cord. Oriental, 12.6 km SE ofLatacunga
Jigeno, Laguna
4000
02°54'S
79° 17'W
Azuay
Cord. Occidental, 13.6 km E ofChaucha
Jilinqui, Cerro
4191
ÜÜ 0 58'S
78°28'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 7.5 km SE ofLaguna de Los Anteojos
o
00
s;: tri s;: o
~ (/)
o ..., :r:tri "rj
Jora-Cruz, Pmo. de
See Pmo. de Y oracruz
Josefo Chiquito, Cerro
4112
01 °07'S
78°43'W
Cotopaxi
Cord. Occidental, 6.4 km SW ofCusubamba
Junacocha,Laguna
4040
02°23'S
78°36'W
Cañar
Cord. Oriental, 9.4 km NW ofHuangra
Ladrillo, Cerro
4160
00° IO'S
78°33'W
Pichincha
8 km NE ofLloa, E ofQuito
Lagoa Toa, Cerro
4180
00°57'S
78°28'W
Cotopaxi
Cord. Oriental, 15.8 km SE·ofLatacunga
o
Lagunas, Pmo. de Las
ztri
~
>-<:
?::1
00°41'N
1rs2w
Carchi
Nudo de Los Pastos, headwaters ofRío Bobo and Río Carchi
7'
o ...,
Laigua Tingo, Cerro
3880
01 °09'S
78°44'W
Cotopaxi
Cord. Occidental, 10 km SW ofCusubamba
Langasa, Cerro
3800
00°49'S
78°25'W
Cotopaxi
Cord. Oriental, 16.6 km S ofVolcán Cotopaxi
Langaya, Cerro
4080
01 o 41'S
78°55'W
Bolívar
Cord. Occidental, 8.8 km E of Santiago
Languachupa, Cerro
4000
ÜÜ 0 51'S
78°2Ü'W
Napo
Cord. Oriental (Cord. de Llanganates), 13.5 km NE ofLaguna de Los Anteojos
Languapungo,Cerro
4040
00°53'S
78°25'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 8 km NW ofLaguna de Los Anteojos
Lanlaguso, Cerro
4295
Ol 0 57'S
78°48'W
Chimborazo
Cord. Occidental, SW ofGuamote
Lanza, Cerro La
4100
01 °25'S
78°48'W
Tungurahua
Cord. Occidental, 4 km NE Nevado Chimborazo
Laurean, Cerro
3460
02°31'S
78°46'W
Cañar
11 km NE oflngapirca
Laurel, Cerro
4000
00°37'S
78°17'W
Napa/Pichincha
Cord. Oriental, between Volcán Cotopaxi and Cerro Antisana, SE ofCerro Sincholagua
León, Pmo. de
4000
Leonan, Cerro ( 1)
4416
01 o 43'S
78°3l'W
Chimborazo
Leonan, Cerro (2)
4000
02°06'S
78°38'W
Chimborazo
10 km SE ofPalmira
Liglepungo, Cerro
3680
02°30'S
78°45'W
Cañar
13 km NE oflngapirca
Limbo, Cerro El
3760
01 °27'S
78°58'W
Bolívar
Cord. Occidental, 17 km W ofVolcán Chimborazo
OJ
:> ~
(")
:>
l'
a
:>
§ ztri
Cotopaxi Cord. Oriental, 7.4 km E ofChambo
<o r-'
00
""'
Elev. (m)
Latitude
Longitud e
Division 1
Notes
Limoturo, Cerro
3600
01 °18'S
78°21'W
Tungurahua
Cord. Oriental, 5 km ESE ofEl Triunfo
Limpiopungo, Pmos. de
3880
00°37'S
78°28'W
Pichincha
Cord. Oriental, Volcán Cotopaxi, NW slopes, around Laguna de Limpiopungo; also called Laguna de Limpias
Lirio, Cerro
4200
01 o 38'S
78°53'W
Chimborazo
Cord. Occidental, 12 km SW of San Juan
Lirio, Cerro El
3800
01 °57'S
78°46'W
Chimborazo
Cord. Occidental, 20 km NE ofPallatanga
Llachapa, Loma
4560
01 °06'S
78°51'W
Cotopaxi
Cord. Occidental, NE of Angamarca
Llactaquingre, Cerro
4000
02°27'S
78°48'W
Cañar
18.8 km NE of Juncal
Lladán, Cerro
3400
02°32'S
78°46'W
Cañar
11 km E oflngapirca
Tungurahua
Uncertain iftrue páramos
N ame
Llangagua, Pmos. de
'-0 '-0
..:::
See Cerro Hermoso
Llanganati, Cerro Llanganati, Pmos. de
3900
01 °09'S
78°18'W
Llano de Sambipamba
3400
00°14'S
78°00'W
Napo
Cord. Oriental, near Oyacachi
Llifatán, Cerro
3800
Ol 0 13'S
78°44'W
Tungurahua
Cord. Occidental, 4.2 km N ofSan Fernando; also spelled Ilifatán
Llilla Cruz, Cerro
4000
02°22'S
78°53'W
Chimborazo
6 km SE ofCompud
Llillán Pungu, Cerro
3480
02°34'S
78°46'W
Cañar
18 km SE ofCañar
Llillín, Cerro
3640
01 °36'S
78°49'W
Chimborazo
Cord. Occidental, 4.5 km NW ofSan Juan
Llimpi, Cerro
3700
01 °22'S
78°34'W
Tungurahua
3.7 km E ofQuero
Llipimachay, Cerro
4200
02°24'S
78°54'W
Chimborazo
11.7 km NE ofJuncal
Lloraquingue, Cerro
4160
02 o 19'S
78°45'W
Chimborazo
Cord. Oriental, 4.6 km S of Achupallas
Lluchir, Cerro
3840
03 °04'S
79°16'W
Azuay
Cord. Occidental, 8.6 km NW of San Fernando
L]ugllugpungu, Cerro
4157
Ol 0 15'S
78°48'W
Tungurahua
Cord. Occidental, 17.7 km NE of Simiatug
Llullucha, Cerro
4025
01 o 35'S
78°48'W
Chimborazo
Cord. Occidental, 5.5 km NW of San Juan
Llullín, Cerro
3622
02°35'S
78°49'W
Cañar
13.2 km SE Cañar
Lluragrumi, Cerro
3920
03 °04'S
79°17'W
Azuay
Cord. Occidental, 9.2 km NW ofSan Fernando
Loca Huañuna, Cerro
4000
00°52'S
78°29'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 14 km NW ofLaguna de Los Anteojos
Loma Gorda, Pmo. de
3850
00°37'S
78° IO'W
Napo
Cord. Oriental, along Río Antisana below Micacocha
Loma Yanaurcu, Cerro
3800
00°08'S
78°34'W
Pichincha
Cord. Occidental, 8 km S ofNono
Luchaloma, Cerro
4320
01 °24'S
78°55'W
Bolívar
Cord. Occidental, 14 km NW ofNevado Chimborazo
Luspa, Pmo. de Laguna
4400
02°46'S
79°!0'W
Cuenca
Cord. Occidental, Cajas area, 44 km W ofCuenca
Lutococha, Laguna
3400
01 °29'S
78°59'W
Bolívar
Cord. Occidental, 20 km SW ofNevado Chimborazo
Machachi, Pmo.
3660
00°08'S
78°05'W
Pichincha
Cord. Oriental, slopes ofVolcán Cotopaxi
Napo/Cotopaxi/Tungurahua
Cord. Oriental (Cord. de Los Llanganates), SE of Chosa Aucacocha, between Aucacocha and Pan de Azúcar
"" > ~ ~ o (/)
N
o
'-0
N
N ame Macho Urcu, Cerro
Elev. (m) Latitude
Longitude
Division 1
Notes
78°48'W
Cañar
18 km NE of Juncal
4000
or26'S
Magtayán, Laguna
3800
02° 14'S
78°35'W
Chimborazo
Cord. Oriental, 11 km E ofTotoras
Malenda Chica, Laguna
3760
01 °23'S
78°4l'W
Tungurahua
4.4 km NW ofMocha
Malenda Grande, Laguna
3763
01 o23'S
78°4l'W
Tungurahua
3.6 km NW ofMocha
Malpán, Cerro
3880
02°18'S
78°51'W
Chimborazo
5 km SW ofGonzol
Mama Tablón, Cerro
3640
02°5l'S
79°07'W
Azuay
5 km NW of Sayausí
Mamarumi, Cerro
4100
00°57'S
79°48'W
Cotopaxi
Cord. Occidental, 1O km W of Zumbagua
Mandana, Cerro
4000
OZ008'S
78°38'W
Chimborazo
12.6 km SE ofPalmira
Mangan, Cerro
3920
02°26'S
78°49'W
Cañar
16 km NE of Juncal
Mangan,Laguna(l)
4000
02°20'S
78°36'W
Cañar
Cord. Oriental, 14.3 km NE ofHuangra
Mangan, Laguna (2)
3840
02°l4'S
78°34'W
Chimborazo
Cord. Oriental, 11.4 km E of Totoras, near Laguna Magtayan
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Mangapungu, Cerro
4240
01 °24'S
78°56'W
Bolívar
Cord. Occidental, 15 km NW ofNevado Chimborazo
Mapahuina, Cerro
4366
02°22'S
78°47'W
Chimborazo/Cañar
Cord. Oriental, SE of Chunchi ·
Marco Quinroy, Laguna
3400
00°26'S
78°20'W
Pichincha
Cord. Oriental, 8.2 km SE ofPintag
Marcos Cocha, Laguna
3600
01 oül'S
78°l7'W
Napo
Cord. Oriental, 23 kni NE ofS~ José de Poaló
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Margarita, Laguna
3440
04° 17'S
79°06'W
Loja
12.5 km SE ofVilcabamba
Mariano Acosta, Pmo. de
3650
00°20'N
78°00'W
Imbabura
Cord. Oriental, along Mariano Acosta-Yahuarcocha rd. (ca. km 20--26)
Mashuo, Cerro
4280
02°24'S
78°49'W
Cañar
17 km NE of Juncal
Matanga,Pmos. de
3500
03 °Ü8'S
78°40'W
Azuay/Morona-Santiago
Cord. Oriental, SSE ofCuenca, SE ofSigsig
Mechahuasca, Pmo. de
4260
01 o 24'S
78°49'W
Tungurahua
Cord. Occidental, N ofVolcán Chimborazo and SW ofCarihuairazo
ñ
Merced, Pmos. La
3100
ÜÜ 22'S
78°20'W
Tungurahua
Cord. Oriental, ENE ofBaños; not true páramo
"Cl
0
Mica, Laguna de La
See Laguna Micacocha 78° 12'W
Napo
Cord. Oriental, in páramo on S slope of Cerro Antisana; also spelled Mica-cocha; also called Laguna de La Mica
ÜÜ 55'S
78°44'W
Cotopaxi
Cord. Occidental, 6.2 km NW ofPujilí
4180
01 o 33'S
78°53'W
Bolívar
Cord. Occidental, Cerro Milín, 12 km NE ofGuaranda
Milín, Pmo. de (2)
4400
01 °Ül'S
78°54'W
Cotopaxi
Cord. Occidental, Pujili-Quevedo rd., NE of Angamarca
Minas, Cerro (1)
4005
00°56'S
78°27'W
Cotopaxi
Cord. Oriental (Cord. de Llanganates), 8 km NW ofLaguna de Los Anteojos
Minas, Cerro (2)
4096
03 °05'S
79°23'W
Azuay
Cord. Occidental, SW of Cuenca, WNW of Girón
Minas, Cerro (3)
4415
02°23'S
78°5l'W
Chimborazo
10 km SE ofCompud
Minas, Cerro (4)
4120
02°10'S
78°39'W
Chimborazo
Cord. Oriental, 9 km NE ofTotoras
Micacocha, Laguna
4100
00°33'S
Milín Chico, Cerro
3810
Milín, Pmo. de (1)
0
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z~
~
~
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00
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N ame _ Minas, Pmo. de
Elev. (m)
Latitud e
Longitude
Division 1
Notes
3600
00° 13'S
78°!2'W
Pichincha
Cord. Oriental, S ofCerro Puntas, NNW ofPapallacta; coordinates estimated
Mintanga, Cerro
3630
00°47'S
78°56'W
Cotopaxi
Cord. Occidental, 2 km W ofChugchilán
Mintza, Pmo. de
4000
O1o27'S
78°20'W
Tungurahua
Cord. Oriental, SE ofVolcán Tungurahua
Mirador, Cerro
3800
01 °05'S
78°44'W
Cotopaxi
Cord. Occidental, 5.5 km SW ofCusubamba
Mirador, Pmo.
4082
00°36'N
7r39'W
Carchi/Sucumbíos
Cord. Oriental, 25 km S ofTulcán, close to Playón de San Francisco; also called Cerro (El) Mirador
Miraflores, Cerro
3400
01 o51'S
78°44'W
Chimborazo
Cord. Occidental, 3 km NW ofColumbe
Miranda, Cerro El
3600
02° li'S
78°55'W
Chimborazo
Cord. Occidental, 5 km NW of Sibambe
Mishquipungincho, Cerro
4359
01 °02'S 01 o24'S
78°49'W
Cotopaxi
Cord. Occidental, 15 km NE of Angamarca
78°45'W
Tungurahua
Cord. Occidental, on Cerro Carihuairazo (5028 m)
00°07'N
78°15'W
Pichincha/Jmbabura
Cord. Occidental (Nudo de Mojanda), S Otavalo Cord. Occidental, 12.6 km NE ofSimiatug
Minza Chica, Pmo. de
May be part ofPmo. de Mintza; more properly spelled Mintza
Mocha Alto, Pmo. Mojanda, Pmos. de
'O 'O 'O
~
4275
Molino Cocha, Laguna
3800
01 °l4'S
78°5l'W
Tungurahua
Molobog Grande, Cerro
3848
02°37'S
78°5l'W
Cañar
11.6 km SE ofCañar
Molón, Cerro
3419
03 °08'S
78°47'W
Azuay
8.5 km S of Cuchil
Monjas, Pmo. de
3915
00°43'S
78°4l'W
Cotopaxi
Cord. Occidental, NW ofToacazo
Montecashra, Cerro
4000
78°44'W
Cotopaxi
Cord. Occidental, 6.4 km SW ofCusubamba
Montero-Urcu, Pmo. de
3600
01 °06'S ooo 12'S
78°12'W
Pichincha
Cord. Oriental, near San Francisco; also spelled Montera-urcu
03 o21'S
79°00'W
Azuay/Morona-Santiago
Cord. Oriental, E ofNabón, SW ofPmos. de Amorgeo
Morochos, Pmos. de
4000
00°20'N
78°20'W
Imbabura
Cord. Occidental, Nevado Cotacachi, SSE slopes, E of Pmo. de Cungas; also spelled Morocho
Mortiños, Cerro
3720
01 o47'S
78°52'W
Chimborazo
15 km SW ofSicalpa
Motero, Pmo. de
3200
Muchanarumi, Pmo. de
3780
Muenala, Pmo. de
4115
00°18'N
78°26'W
Imbabura
Cord. Occidental, W ofVolcán Cotacachi; possibly only 3631 m
Muerte Pungo, Laguna
4000
00°25'S
78°16'W
Pichincha
Cord. Oriental, 12.7 km SW ofPintag
01 °5l'S 01 o16'S
78°52'W
Chimborazo
Cord. Occidental, 17.3 km NE ofPallatanga
78°45'W
Tungurahua
Cord. Occidental, 22.3 km NE ofSimiatug
Cotopaxi
Cord. Oriental, part ofPmos. de Pisayambo
Morire, Pmo. de
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Pichincha Pichincha
Cord. Oriental, Cerro Sincholagua, E slopes
Mulachimbana, Cerro
3800
Mulanleo, Cerro
3400
Mulatos, Pmo. de
4100
Mullos, Cerro
3856
01 °57'S
78°53'W
Chimborazo
Cord. Occidental, 10 km NE ofPallatanga
Mulmul, Cerro
3800
01 °25'S
78°32'W
Tungurahua
4 km W ofCotaló
Muru Urcu, Pmo.
3950
00°47'S
78°24'W
Cotopaxi
N
N ame
Elev. (m)
Latitude
Longitude
Naba, Pmo. de Nabón, Pmo. de
3600
Division 1
Notes
Chimborazo
Old name used by Spruce
Azuay
Old name used by Jameson; perhaps refers to the Tinajilla area since tbe town of Nabón is only at 2750 m
N
Ñ
Naranjito, Cerro
3800
01 a54'S
78°47'W
Chimborazo
Cord. Occidental, 21.8 km NE ofPallatanga
Narhuiña, Cerro
3904
03 °09'S
79°32'W
Azuay
W ofGirón, NW ofPucará; also spelled Narihuiña
Nariz del Diablo, Cerro
3800
OOa 12'S
78°35'W
Pichincha
Cord. Occidental, 5.5 km NW ofL!oa, W ofQuito
Natosa, Laguna
3640
04°43'S
79°25'W
Loja
5 km SW ofLaguna Cox
Nauna, Cerro
3560
02°ll'S
78°48'W
Chimborazo
Cord. Oriental, 3.6 km NE of Alausí
Naupán, Cerro
4280
78°5l'W
Cañar
Cord. Oriental, 11 km SE ofCompud
Navag, Pmos. de
3900
02°23'S 01 a46'S
78°50'W
Chimborazo
SW ofSicalpa, W ofPmos. de Quillotoro, SE ofPmos. de Huangupud
Negra, Laguna(!)
4000
02°50'S
79° 12'W
Azuay
Cord. Occidental, 15 km NW ofSayausí
Negra, Laguna (2)
3450
04 a45'S
79°25'W
Loja
7.5 km SW ofLaguna Cox
::r:
Negra, Laguna (3)
3800
00°08'S
78°l5'W
Pichincha
10.3 km NE ofTocachi
ztTJ
Negra, Laguna (4)
4480
01 °24'S
78°46'W
Tungurahua
Cord. Occidental, 7.4 km NE ofNevado Chimborazo
Negra, Pmo. de la Laguna
4100
02° IO'S
Chimborazo/Morona-Santiago
Cord. Oriental, 20.7 km NE-o.fTotoras, NE of Alao
Negro Puñona, Cerro
4000
01 °08'S
78°30'W 78a43'W
Cotopaxi
Cord. Occidental, 8.2 km SE o(Cusubamba
03°0l'S
79°22'W
Azuay
Cord. Occidental, 19 km NW ofSan Fernando
See Cerro Narhuiña
Narihuiña, Cerro
Negro, Cerro (1)
4080
Negro, Cerro (2)
4460
Negro, Cerro (3)
4200
Negro, Cerro (4) N egros, Cerros
~
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tTJ
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Carchi 0
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78°3l'W
Chimborazo
Cord. Oriental, 7 km NE of Chambo
3786
Ol 42'S 03 a35'S
79°23'W
Loja
7 km W ofCelén
3520
04°3l'S
79°l8'W
Zamora-Chinchipe
14.3 km NE of Amaluza
Nignig, Cerro
3720
00°49'S
78°45'W
Cotopaxi
15 km SE oflsinliví
Ninaguilca Chico, Cerro
3600
00°22'S
78°38'W
Pichincha
11.2 km NW ofTambillo
Ninaguilca Grande, Cerro
3788
00°22'S
78°40'W
Pichincha
Cord. Occidental, 14 km NW of Alóag Cord. Oriental, 4. 7 km SW of San José de Poaló
Niño Urcu, Cerro
3425
00°53'S
78°42'W
Cotopaxi
Niñococha,Laguna
3600
02°23'S
78°33'W
Cañar
Cord. Oriental, 10 km NE ofHuangra
Niñoloma, Cerro
3680
00° 19'S
78°35'W
Pichincha
Cord. Occidental, 7 km SW ofChillogallo, SW ofQuito
Novillo Pungu, Cerro(!)
4455
01 °06'S
78°49'W
Cotopaxi
Cord. Occidental, 12 km NE of Angamarca
Novillo Pungu, Cerro (2)
4018
01 °09'S
Tungurahua
Cord. Occidental, 10 km S ofCusubamba
Ñuctushca, Cerro
3480
02°35'S
78°41'W 78a46'W
Cañar
18 km SE ofCañar
z
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00
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N ame
Elev. (m) Latitude
Longitude
Division 1
Notes Cord. Oriental, 18 km SW ofOyacachi; sometimes misspelled Numalviro
00° 15'S
78°12'W
Pichincha
Nutorrumi, Cerro
3820
03°0l'S
79°!0'W
Azuay
13 km NW ofVictoria del Portete
Oreja, Cerro La
4400
7r58'W
Carchi
Nudo de Los Pastos, near Colombia border, W ofTufiño
. Nunalviro, Laguna
Orquetacocha, Laguna
4260
00°49'N 01 o2l'S
78°44'W
Tungurahua
Cord. Occidental, 9.3 km SW ofTisaleo
Osobolsa, Cerro
3400
01 o43'S
78°57'W
Bolívar
Cord. Occidental, 6 km SE ofSantiago
Osogochi, Pmo.
3780
02°14'S
78°36'W
Chimborazo
Cord. Oriental, 7 km NE ofTotoras
Osotambo, Cerro
3720
01 °04'S
78°l8'W
Napo
Cord. Oriental, 18 km NE of San José de Poaló
Napo
Osthana, Pmo. Oyacachi, Pmos. de
Pichincha/Napa 00°42'S
Pachusala, Pmo. de
78°4l'W
Cord. Oriental, general name for páramos in vicinity of Oyacachi
Cotopaxi
Padre Encantado, Cerro
4600
00°09'S
78°34'W
Pichincha
Cord. Occidental, part ofMacizo del Pichincha, SW ofQuito
Padre Rumi, Cerro
4330
00°49'S
78°48'W
Cotopaxi
SE of Isinlibí
Paja Blanca, Pmo. de
3300
Palama, Pmo.
Carchi
Cord. Oriental, interandean area
Cotopaxi
Cord. Oriental, W slopes, E of(San Miguel de) Salcedo
Chimborazo
Between Guamote and Alausí
Palmira, Pmo. de
3229
02°05'S
78°45'W
Palmurco, Pmos. de
3200
03°l6'S
78°56'W
Azuay
17.2 km on Jima-San Miguel de Cuyes rd.
Pamba-Marca, Cerro
4075
00°07'S
78°l2'W
Pichincha
ESE ofGuayllabamba, ENE ofEl Quinche; also spelled Pambamarca
Pan de Azúcar, Cerro
4000
01 °08'S
78°18'W
Cotopaxi
Cord. Oriental, 20 km from San José de Poaló
"O
> ;;x:l > ;;: o (l:l
Chimborazo
Pangor, Pmo. de Pansache,Pmo. de Panza, Pmo. de
::0 \.0 ..::;
4275
Cotopaxi
Also spelled Panzache
Chimborazo
Cord. Occidental, Volcán Chimborazo, S slopes See Pmo. de Guamaní
Papallacta, Pmo. Páramo Chico, Loma
3800
Paredones, Pmo. de
4085
Pasa, Pmo. de
4000
Pasochoa, Pmos. del
4199
00°06'S
78°08'W
Pichincha
Cord. Oriental, SE ofCangahua
Chimborazo
S ofRiobamba
Cotopaxi 00°25'S
78°29'W
Pichincha
Cord. Oriental, slopes ofVolcán Pasochoa, SE ofTambillo, S of Amaguaña; also spelled Pasuchoa; name derived from Paso Ochoa (F. Sarmeinto, pers. comm.)
Azuay
Cord. Oriental, between Gualaceo and Limón
Patococha, Pmo.
3450
Patul, Cerro
4168
02°4l'S
79°04'W
Cañar
Cord. Occidental, NW of Azogues, W ofBiblián
Pedregal, Pmo. del
3600
00°30'S
78°29'W
Pichincha
Cord. Oriental, NW of Sincholagua
Peila, Cerro
4473
02°26'S
78°47'W
Cañar
NE ofTambo N
w
N
N ame
Elev. (m)
Latitude
Longitude
Division 1
Notes
Pelado, Cerro El
4151
00°44'N
7r54'W
Carchi
25 km SW ofTulcán
Picachos, Cerro Los
3720
04 o 42'S
79°24'W
Loja
3 km SW ofLaguna Cox
Pichincha, Pmo. del
4000
00°10'S
78°34'W
Pichincha
Cord. Occidental, slopes ofCerro Rucu Pichincha and Guagua Pichincha, W of Quito
Piedramagna, Cerro
4432
01 °30'S
78°38'W
Tungurahua!Chimborazo
Cord. Occidental, E ofVolcán Chimborazo, N ofRiobamba Cf. Acosta-Solís, 1984
Pilis-urcu, Pmo.
See Pmo. de Piñán
Piñal, Pmo. de Piñán, Pmo. de
4536
00°30'N
78°35'W
Imbabura
Cord. Occidental (incl. Cerro Yana-Urcu), NW oflbarra, W ofTumbabiro, N of Volcán Cotacachi; also called Pmo. de Piñal
Pinlilligue, Pmo. de
3700
01 °52'S
78°27'W
Chimborazo
Cord. Oriental, between Alao and La Tranca See Pmo. de Pinlilligue
Pinllillic, Pmo. de Pisayambo, Pmos. de
3900
01 °06'S
78°22'W
Cotopaxi
Cord. Oriental, ENE ofSan José de Poaló; incl. Laguna(de) Pisayambo, which is just across the border in Tungurahua
Plazuela, Cerro La
4320
00° lü'S
78°34'W
Pichincha
Cord. Occidental, 8 km N ofL!oa
01 °23'S
78°43'W
Tungurahua
Cord. Occidental, SW ofTisaleo, NW ofMocha
Pichincha
Cord. Occidental, Cerro El Corazón, N slopes, close to Hacienda La Granja; possibly equals Pmo. Grande. (2)
79°16'W
Azuay
Nudo de Portete Cord. Oriental (Cord. de Llanganates), 8 km NE ofLaguna de Los Anteojos
Pocacocha, Cerro
4200
Pongo, Pmo. El
3450
Portete, Pmo.
3600
03 °05'S
Preñado, Cerro El
4043
00°53'S
78°23'W
Napo
Pucará, Pmo. de
3600
00° 17'S
78°17'W
Pichincha
Puebolo, Volcán
3950
Cord. Oriental, summit ofPucará is 3600 m Cord. Occidental, W ofVolcán Chimborazo, along Zanjapamba-Simiatug rd.; an extinct volcano (Acosta-Solís, 1984: 76) Cord. Occidental, W ofVo1cán Chimborazo, at base ofVolcán Puebo1o
Puenebata, Pmo. de Pujilí, Pmo. de
3800
00°54'S
78°43'W
Cotopaxi
Cord. Occidental, W ofLatacunga toward Zumbahua
Púlpito, Cerro El (1)
3927
01°14'S
78°27'W
Tungurahua
Cord. Oriental, 4 km NE of Sucre
Púlpito, Cerro El (2)
4200
02°17'S
78°34'W
Chimborazo
Cord. Oriental, 12.5 km SE ofTotoras
Pumachaca, Cerro
4260
02 a 17'S
78°49'W
Chimborazo
E ofChunchi, SE ofSibambe
S:: tn S::
o
~
Cll
o ..., ::r: tn "rj
ztn
:E
-< o ~ til o..., z> ñ > L'
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§ ztn
Chimborazo (?)
Pungapala, Pmo. de Puntas, Cerro
4452
00°l!'S
78°!2'W
Napo/Pichincha
Cord. Oriental, Estribuciones de Oyacachi, 33 km E of Quito
Puruhanta, Laguna
3750
00°12'N
77°57'W
lmbabura
Cord. Occidental, ENE ofVolcán Cayambe; also spelled Puruhanta; also called Laguna Chique or Laguna de Chiquí
4000
01 o 42'S
78°52'W
Putzalagua, Pmo. Puya!, Pmo. de
_¡,.
Cotopaxi
Cord. Oriental, NE of (San Miguel de) Salcedo
Bolívar
Cord'. Occidental, slopes ofCerro Puya! (4373 m), W ofCajabamba, E ofGuaranda
<o r
00
_¡,.
N ame
Elev. (m)
Latitud e
Longitude
Division 1
Qinualoma, Cerro Quilimás, Pmo.
Notes See Cerro Quinaloma
4710
Chimborazo
Cord. Oriental, slopes ofVolcán Quilimás, SE ofRiobamba; also called Cubillín and possibly Pmo. de Cubillín (Diels, 1937: 55); misspelled Cubullín
Quilindaña, Pmo.
4877
00°45'S
78°19'W
Napo
Cord. Oriental, 18 km SE ofVolcán Cotopaxi (near Laguna Yurac Cocha)
Quillotoro, Pmos. de
4051
01 o46'S
78°49'W
Chimborzao
SW ofSicalpa, E ofPmos. de Navag
Quillu Urcu, Cerro
4572
01 °04'S
78°52'W
Cotopaxi
Cord. Occidental, SW of Latacunga, NW of Angamarca
Quilotoa, Volcán
3981
78°55'W
Cotopaxi
Cord. Occidental, 32 km W ofLatacunga
Quimsacocha, Pmo. de
3969
00°52'S 03 ooo·s
79°l5'W
Azuay
Cord. Occidental, NW ofGirón, N ofSan Fernando
Quinaloma, Cerro
3934
02°29'S
78°40'W
Cañar/Morona-Santiago
Cord. Oriental, SE ofChunchi, ENE ofCañar; also spelled Qinualoma
Quingo, Cerro El
3680
01 °16'S
78°47'W
Tungurahua
Cord. Occidental, 19.3 km NE ofSimiatug
Quinoas, Pmo.
3600
Azuay
Cord. Occidental, W of Cuenca
QuinsaCruz
4445
Cañar
S ofCañar
Quispicacha, Cerro
4585
01 °07'S
78°46'W
Tungurahua
Cord. Occidental, SW of San Miguel de Salcedo
Redondo, Cerro El
3835
00°22'N
78°04'W
Imbabura
Cord. Oriental, SE of !barra, E of Caranqui
01 °35'S
78°46'W
Chimborazo
Resgón, Pmo. de Retanales, Pmo. de
3400
Reventador, Pmos. del
3483
00°03'S 01 °07'S
"'::!
;¡:..
Cotopaxi
Vicinity ofLatacunga and Laguna Salayambo
77°40'W
Napo
Cord. Oriental, part ofVolcán Reventador
78°52'W
Cotopaxi
Cord. Occidental, E of Angamarca
Ricuri, Cerro
4360
Romerillos, Pmo. de
3400
Cotopaxi
Romo, Pmo.
3700
Tungurahua/N apo
Rucu Pichincha, Pmo.
4500
00°10'S
78°34'W
Pichincha
Cord. Occidental, NE side ofVolcán Rucu Pichincha
Rumiñahui, Cerro
4722
00°34'S
78°30'W
Pichincha!Cotopaxi
Cord. Oriental, 15 km NW ofVolcán Cotopaxi Cord. Oriental, slopes ofCerro Rumiñahui
~ s;:: oC/J
Cord. Oriental (Cord. de Los Llanganates), E of San José de Poaló, between Sinchilibí and Río Potrero
Rumiñahui, Pmos. de
4719
Pichincha
Rumipungu, Pmo.
3610
Cotopaxi
Cord. Oriental, slopes ofCerro Quilindaña
Sacraurcu, Cerro
4545
02°03'S
Chimborazo
ENE ofPalmira
78°33'W
'D 'D
~
Sagatoa, Cerro
4152
01°12'S
78°35'W
Cotopaxi/Tungurahua
Cord. Occidental, W ofPíllaro; also spelled Sagoatoa
Salasacas, Pmo. de
3860
01 o24'S
78°45'W
Tungurahua
Cord. Occidental, on Cerro Pocacochas, 2 km from the E glacier ofVolcán Carihuairazo
Salayambo, Pmo. de Laguna
4100
Cotopaxi
Cord. Oriental, E ofLatacunga; also spelled Salayampe
Salcedo, Pmos. de
3850
00°55'S
78°28'W
Cotopaxi/Napo
Cord. Oriental, páramos E ofSalcedo, collectively also known as the Langoa (F. Sarmiento, pers. comm.)
N V,
N ame
Elev. (m) Latitude
Salto, Cerro El
3800
Salto, El
4160
02°02'S
Longitud e Division 1 78°5l'W
N
Notes
Chimborazo
Cord. Occidental, 13 km NW ofTixán
Carchi
Nudo de Los Pastos, 0.5 km N ofCerro El Pelado
San Agustín, Pmo.
3400
00°41'S
78°32'W
Cotopaxi
Cord. Oriental, Volcán Cotopaxi, SW slopes
San Antonio, Cerro
4000
04°45'S
78°23'W
Zamora-Chinchipe
S ofNudo de Sabanilla
San Elías, Pmos.
3300
00°45'S
78°32'W
Cotopaxi
Cord. Oriental, NE ofMulaló
San Francisco, Cerro de
4685
San Juan, Pmo. de
Carchi
Cord. Oriental, also known as Playón de San Francisco
Pichincha
Cord. Occidental, probably the area around San Juan de Chiriboga above Quito along old rd. before descent to Santo Domingo
San Marcos, Pmo. de
3360
00°07'N
77°52'W
Pichincha
Cord. Oriental, N slopes of Volcán Cayambe; incL Laguna San Marcos more properly known as Pishcococha (F. Sarmiento, pers. comm.)
San Vicente, Cerro
3710
02°35'S
78°37'W
Cañar
Cord. Oriental, NNE ofRivera; páramo?
Sanancajal, Pmos.
3610
00° 10'S
78°55'W
Chimborazo
Cord. Occidental, Volcán Chimborazo, E slopes; spelled Sanancajas by Whymper (1882)
Sanancajas, Pmo. de
See Pmos. Sanancajal
Sangay, Volcán
5230
01 o58'S
78°22'W
Morona-Santiago
Cord. Oriental
Santa Rosa, Cerro
4000
01 °24'S
78°43'W
Tungurahua
Cord. Occidental, SSW ofT~saleo
Sara-Urcu, Cerro
4676
00007'S
77°55'W
Pichincha!Napo
Saraguro, Pmo. de
3300
Saratano, Cerro El
4100
Sebritana, Pmo. de
3400
Shinigually, Cerro
Cord. Oriental, NE ofVolcán Antisana, S ofVolcán Cayambe; also spelled Saraurcu Close to Oña
00°34'N
77°37'W
4198
02°05'S
78o38'W
Shorshay, Cerro
3962
02°23'S
78°40'W
Siguata, Pmos. de
4000
Silla, Cerro La
3980
02°27'S
Sillacajas, Cerro
4211
02°l2'S
Silván, Pmo. de
3200
Azuay
37 km S ofCuenca
Simiatug, Pmo. de
3500
Bolívar
Cord. Occidental
Sincholagua, Pmos. de
4900
Pichincha
Cord. Oriental, slopes ofCerro Sincholagua, between Volcán Antisana and Volcán Cotopaxi
Carchi
Cord. Oriental (Cord. de Mirador), S ofEl Playón de San Francisco
Pichincha
Cord. Oriental, E slopes of Cotopaxi, near Volcán Sangay
Chimborazo
SE ofGuamote, E ofPalmira
Chimborazo/Cañar/MoronaSantiago
Cord. Oriental, ESE ofChunchi
Cotopaxi
Cord. Occidental, Cerro Illiniza, N slopes
78°49'W
Cañar
15.8 km NE of Juncal
78°42'W
Chimborazo
Cord. Oriental, E of Alausí
00°32'S
78°23'W
Soguillas, Pmo. de
4000
01 °08'S
78° 15'W
Napo
Cord. Oriental, near Las Torres de Llanganati
Soldados, Pmo. de
4138
02°54'S
79°20'W
Azuay
Cord. Occidental, Cerro Soldados, W ofCuenca
o;
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o
~
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o'Tj >-3 ~
tTJ
ztTJ
:E --< o
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to
o >-3 >z ñ >r
a >-
§ tTJ z
<o r-'
00
_¡:,.
N ame
Elev. (m)
Latitude
Longitude
Division 1
Notes
Sombrero Urcu, Cerro
4511
01 °!2'S
78°52'W
Bolívar
Cord. Occidental, 15 km NE ofSimiatug
Soroche, (Cerro ?)
4730
or2o·s
78°32'W
Cañar/MoronaSantiago
Cord. Oriental, W ofMacas, SE ofLaguna de Ozogoche, Laguna Aucacocha is on its N slopes
Sumaco, Volcán
4092
00°34'S
7r38'W
Napo
Cord. Oriental or sometimes called the Tercera Cordillera (Cord. de los Huacamayos)
Tablachupa, Pmos. de
4000
78°23'W
Imbabura
Cord. Occidental, SW ofNevado Cotacachi, W ofPmo. de Cungas
Tablón, Cerro El (1)
3880
00°20'N 01 o 24'S
78°59'W
Bolívar
Cord. Occidental, 20 km NW ofNevado Chimborazo
Tablón, Cerro El (2)
3800
00°23'N
78°20'W
lmbabura
Cord. Occidental, 17.4 km NW of Cotacachi
Tilila, Pmo. de
3500
Bolívar
Cord. Occidental
Tililac, Pmo. de
4000
Chimborazo
Cord. Occidental, above San Juan, along rd. to Guaranda; sometimes misspelled Tolallac
Tinajillas, Pmo. de
3500
Cord. Oriental, 30-50 km S ofCuenca
Tingo, Pmo. el Tioloma, Cerro (de)
4263
Tiopullo, Pmo. de
3500
03 o 13'S
79°04'W
Azuay
01 °0l'S
78°48'W
Cotopaxi
02°20'S
78°38'W
Tipococha, Pmo. de
Chimborazo
Nudo de Tiocajas, E ofChunchi
Pichincha/Cotopaxi
Cord. Occidental, near Lloa Chiquito, near Illiniza; also called Nudo de Tiopullo and Pmo. Güintza (Huinsa)
Chimborazo
Nudo de Azuay
Tiuguinal, Cerro
4097
01 o22'S
79°001W
Bolívar
Cord. Occidental, 4 km NNE of Salinas
Toldo, Cerro (1)
4402
01 °44'S
78°30'W
Chimborazo
Cord. Oriental, E ofRiobamba
Toldo, Cerro El (2)
4200
02° 16'S
78°37'W
Chimborazo
Cord. Oriental, 6.7 km SE ofTotoras
Toledo, Cerro
3200
04°23'N
79°07'W
Loja/Zamora
Nudo de Sabanilla, E ofVilcabamba
Toreadora, Pmo. (de la)
3930
Azuay
Cord. Occidental, vicinity ofLaguna Toreadora, between Molleturo and Quinoas
78°44'W
Cañar
Cord. Oriental, 20 km NW ofHuangra
Torre, Cerro La ( 1)
4253
02°20'S
Torre, Cerro La (2)
3720
01 °48'S
78°51'W
Chimborazo
Cord. Occidental, 3 km NE of Juan de Velasco
Totorilias, Pmo. de
4015
01 °30'S
78°49'W
Chimborazo
Cord. Occidental, on Volcán Chimborazo; sometimes misspelled Totovillas
Tres Cruces, Pmo. de Las
3500
Chimborazo/Morona-Santiago
Cord. Oriental, N of Alao
Trujillo, Pmo. de
3450
Pichincha/Cotopaxi
Between peaks Iliniza and Cotopaxi
Tufiño, Pmo. de
3500
Carchi
Nudo de Los Pastos, W ofTulcán, part of greater Pmos. del Angel
Tuja, Pmo. de
3400
Carchi
Also called Pmo. de Tusa, Pmo, de Tunza, and Pmo. de Tuxa
"ti
;¡,. ?:1 ;¡,. ~
o
[/l
See Pmo. de Zumbagua
Tungua, Pmo. de Tungurahua, Volcán
'O 'O
.:S
5005
01 °27'S
78°26'W
Tungurahua
Cord. Oriental, 15 km SSW ofBaños
Tunza, Pmo. de
See Pmo. de Tuja
Tusa, Pmo. de
See Pmo. de Tuja
N
...,
N
Elev. (m)
Latitude
Longitude
Division 1
Tzhigua, Cerro
4246
01 °04'S
78°56'W
Cotopaxi
Ungüi, Cerro
3584
Urbina, Pmo. de
4050
Vaquería, Pmo. de La
4200
N ame
Notes See Pmo. de Tuja; Tuxa is an old name
Tuxa, Pmo. de
Veracruz, Pmo. de
Cord. Occidental, NNW of Angamarca, S ofQuindigua
Pichincha
Cord. Occidental, SW ofQuito
01 °26'S
78°43'W
Chimborazo
Cord. Occidental, Nudo de Igualata Sanancajas, SE ofVolcán Carihuairazo and E ofVolcán Chimborazo Cord. Occidental, NE ofPallatanga, E ofPmo. Huaico
01 a56'S
78°48'W
Chimborazo
00°25'W
78°00'N
Imbabura See Pmo. de Guamaní
Virgen, Pmo. de La Viudita, Cerro La
3786
00°25'S
78°39'W
Pichincha
Cord. Occidental, SW ofQuito, NW ofMachachi
Voladero, Pmo. del
3815
00°4l'N
7r53'W
Carchi
Nudo de Los Pastos, part ofPmo. de El Angel along El Angel-Tulcán rd., actually this is the site of Laguna El Voladero
Volcán, Cerro El
3600
00°09'S
78°39'W
Pichincha
Cord. Occidental, 13.3 km SW ofNono
Yana-Ashpa
4185
01 a42'S
Chimborazo/Bolívar
Cord. Occidental, SE ofGuaranda, E ofSan José
Yana-Urcu, Cerro(!)
4536
00°28'N
78°50'W 78a20'W
Imbabura
Cord. Occidental, NW of !barra,. W of Tumbabiro (incl. Laguna Negra or Yanacocha); also called Cerro Yanahurcu de Piñan
Yana-Urcu, Cerro (2)
;;:;
4116
02°26'S
78°39'W
Cañar
NEofTambo See Cerro Yana-Urcu (!); Yanahurco comes from Quichua base Yana Urcu
Yanahurcu de Piñán, Cerro 01 a04'S
Cotopaxi
Cord. Occidental, 20 km ENE of Angamarca, E ofPmo. de Chilca
Imbabura
Cord. Oriental, along rd. E from Ibarrá; also written as Jora-Cruz
78°05'W
Imbabura
10 km N oflbarra, on Hacienda Yura Cruz, highest point that separates Laguna Yaguarcocha with the Pimampiro drainage towards Río Chota; also as Yuracruz
00°47'S
78°2l'W
Napo
Cord. Oriental, 3 km E of Cerro Quilindaña; páramos around the Iake
03 ooo·s
78°35'W
Azuay
Cord. Oriental, near Gualaceo-Limón rd.
Loja
Between Oña and Saraguro; old name used by Lehmann
Yanaurcu, Pmo. de
4227
Yoracruz, Pmo. de
3000
Yura Cruz, Pmo.
3800
00°22'N
Yuragcocha, Laguna
4050
Zapote Naida, Pmo. de
3400
Zhozunes, Pmo. de
3300
78a46'W
See Pmos. de Zumbahua
Zumbagua, Pmos. de Zumbahua, Pmos. de
4200
Zuñiga, Pmo. de
4091
00°22'S
78°36'W
Cotopaxi
Cord. Occidental, Pilaló-Latacunga rd.; also misspelled as Zumbagua
Pichincha
Cerro Atacazo, S slopes, S ofQuito
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1999]
PARAMOS
219
Peru James L. Luteyn
The páramos of Peru, locally called "jaleas," are located north of 8°S latitude in the northem portions ofthe Cordillera Occidental and the Cordillera Oriental, in the departments ofPiura, Cajamarca, Amazonas, Lambayeque, San Martín, and La Libertad. Peruvian páramos are very similar to those of southem Ecuador in vegetation and general physiognomy. They may begin at the low elevations of2900-31 00 m especial! y east ofthe Río Marañón, where the cordillera is lower in general and moister air from the Amazon basin predominates. The páramos ofPeru seem to be more grass-dominated and less shrubby overall, with many more rock outcrops. In the region north ofthe city of Cajamarca, the páramos are intensively cultivated up to about 3500 m with com and potatoes; between 3500
m and about 3800 m there are now many Pinus plantations. In the more eastem páramos, around the Jaleas de Calla Calla (Balsas-Leimebamba road), for example, there is less population pressure and the páramos may occur at elevations as low as 3000 m. In Peru, páramo (jalea) includes a total of 420,000 hectares (I. Sánchez Vega, pers. comm.) orca. 4200 km2 • The only Peruvian national park within our range that includes páramo is Río Abiseo, although Huascarán (just south of 8°S latitude in the Cordillera Blanca) may also have páramo (cf. D. N. Smith, 1988). For additional discussion about Peruvian páramo vegetation, see D. N. Smith, 1988; Weberbauer, 1911, 1936, 1945; Young & León, 1988, 1990; Young & Reynel, 1997; and Young et al., 1997.
N ame Alaska, Paso de
Elev. (m)
Latitud e
Longitude
4000
Division 1
Division 2
Notes
La Libertad
Pataz
Paseo y-Tayabamba rd.
Algamarca Hualaco
3850
Cajamarca
Cajabamba
N ofRío Huacaday
Alturas de Huallamarca
3580
La Libertad
Pataz
Huaguil-Pallar rd.
3500
Cajamarca
Cajamarca
Near Porcón; also spelled Atashaico
Amazonas
Chachapoyas
07°37'S
78°19'W
o
See Jalea de Atocsaicu
Atashaico, Jalea de Atocsaicu, Jalea de
N N
06°47'S
Barro Negro, Paso
3950
Baúl, Cerro
4140
Calla Calla, Jaleas de
3750
06°47'S
Camich
3400
or15'S
Campanario, Cerro (1)
3900
Campanario, Cerro (2)
3500
7r51'W
Cajamarca
Cajamarca
77°5l'W
Amazonas
Chachapoyas
1rss'W
Cajamarca
Cajabamba
Amazonas
Chachapoyas Cajamarca
oro3'S
78°39'W
Cajamarca
Balsas-Leimebamba rd.; extensive area ofjalea ranging from ca. 3000 to 3750 m
o
NW ofCajamarca, along rd. to Porcón
3700
06°46'S
78°37'W
Cajamarca
Hualgayoc
3 km W ofHualgayoc
4050
or59's
78°29'W
Cajamarca
Cajamarca
S ofLaguna,Yanacocha
Cascabamba, Jalea de
3600
or23'S
78°42'W
Cajamarca
Contumazá
15 km E of Contumazá, a1ong rd. to Saleot
Cerro Negro, Pampa de
4700
oroo·s
78°35'W
Cajamarca
Cajamarca
Km 27 on Cajamarca-Hualgayoc rd.
Cha1huayacu
3550
oro3'S
78°10'W
Cajamarca
Celendín
BetWeen Encañada and Celendín
Challuayaco, Jalea de
3150
Cajamarca
Cajamarca
Chamis, Laguna
3300
oro7's
78°34'W
Cajamarca
Cajamarca
Chimchim
3650
or34'S
78°20'W
Cajamarca
Cajabamba
Chugur
3200
or33'S
78°20'W
Cajamarca
Cajabamba
Chugurpata
3200
or32'S
7rs9'W
Cajamarca
Cajabamba
Chuño, Jalea El
4500
Cajamarca
Contumazá
Also called Pozo Chuño
Cinchao, El
3750
06°43'S
Cajamarca
Hualgayoc
Along rd. to Chugur
Coechan
3550
78°00'W
Amazonas
Luya
Along Luya Viejo-Luya Chico rd.
La Libertad
Santiago de Chuco
See Jalea éle -~oymolache
78°39'W
Coimolache, Jalea de
2:: o
~
Carachugo Sur, Cerro
. 06°09'S
tn
(/l
Candela, Cerro
Caymolache
S:
NW of Cajamarca
"rl ~ """ tn
ztn
~
o-<
~
l:lJ
o ~ """ z ñ
~
r
o ~
~ ~
See Jalea de Coymolache
Coipín, Jalea de
4000
Co llana, Cerro
3814
oso 16'S
79o 18'W
Amazonas? (La Libertad)
Collpa Grande
3600
or3s's
1rs6'W
Cajamarca
Cajabamba
Combayo
3200
oro1·s
78°25'W
Cajamarca
Cajamarca
Distrito de Encañada
<o r
00
_¡,.
Name
Elev. (m)
Latitude
Longitud e Division 1
Division2
Notes See Jalea de Kumulea
Comulca, Jalea de 05°05'S
79°4l'W
Piura
Morropón
3650
07"0l'S
78°23'W
Cajamarca
Cajamarca
3320
78°28'W
06°5l'S
Cajamarca
Cajamarca
Coyma, Cerro
3350
05°56'S
79°26'W
Piura
Huancabamba
Coymolache, Jalea de
3900
06°42'S
78°40'W
Cajamarca
Hualgayoc Huamachuco
Confeccionado
3200
Conor Corral pampa
Km 18 along Cajamarca-Bambamarco rd., W ofHualgayoc; also spelled Coimolache and Caymolache
Cuchi Corral, Cerro
3800
07"52'S
78°02'W
La Libertad
Cuello del Indio
3200
05°20'S
79°32'W
Piura
Huancabamba
Near Surupite, along rd. to Huancabamba
Cueva Negra
3800
La Libertad
Pataz
Near Tuco, Tayabamba-Huancaspata
Culebra, Pampa de la
3200
Cajamarca
Cajamarca
Between Cajamarca and La Encañada
Culquirumi
3300
Cajamarca
Hualgayoc
oro8'S
78°23'W
Cumbe Mayo, Jalea de Cumbemayo, Jalea de
95 km along the Cajamarca-Bambamarca rd. See Jalea de Cumbemayo
3600
07"12'S
78°36'W
Cajamarca
Cajamarca
Cumullka, Jalea de Empalme
'D 'D
2S
3600
06°50'S
78°4l'W
Cajamarca
Hualgayoc
10 km WSW ofCajamarca, along rd. to Chetilla; also spelled Cumbe Mayo
;p.
See Jalea de Kumulca
S:
Cajamarca-Hualgayoc rd.
o
Encajon, Quebrada
3850
07"0l'S
78°32'W
Cajamarca
Cajamarca
W of Minera Y anacocha
Espino, El
3550
07"37'S
78°22'W
Cajamarca
Cajabamba
Near border with the Departamento de La Libertad
Estacas, Jalea de Las
2350
Cajamarca
San Miguel
Fraijaco (Huauui-Huni), Jalea de Cerro de
3450
Amazonas
Gavilán, Jalea de Cerro
3560
Huaguil, Jalea de
4000
Huahuiña, Cerro
3700
Cajamarca
Hualgayoc
Hualgayoc
3900
06°47'S
78°36'W
Cajamarca
Hualgayoc
Huancacarpa
3400
05°08'S
79°32'W
Piura
Huancabamba
07"12'S
78°3l'W
"" 2;:
(l:l
Quisuarpampa (El Tingo area) NE ofTambo de Ventilla
Cajamarca
Cajamarca
Abra El Gavilán, S of Cajamarca, Cajamarca-Chilete rd.; also called El Gavilán
La Libertad
Sánchez Carrión or Huamachuco
Paliar-Molino Viejo rd.; also called Cerro Consuy?
Above Culquirrumi; sometimes spelled Gualgayoc
Huanico
3500
07"09'S
78°08'W
Cajamarca
Celendín
Cajamarca-Celendín rd., toward Huagal
Huayllidas, Señal
4733
07"53'S
78°02'W
La Libertad
Sánchez Carión
Above Lagunas de Huangagocha
J alampa, Jalea de
3800
Cajamarca
Hualgayoc
Km 35 on Cajamarca-Hualgayoc rd.; also spelled Jelampa N N
Elev. (m)
Latitude
Longitude
Jelig, Jalea de
3200
06°53'S
78°07'W
Julia, Pampas de La
3600
N ame
Division 1
Division2
Notes Balsas-Celendín rd.; also called Paso Gelig or spelled Jelic
Cajamarca
Celendín
La Libertad
Santiago de Chuco
Kahuish, Jalea de
3500
Cajamarca
Cajamarca
Along rd. to Celendín
Kumulca, Jalea de
3850
07°05'S
78°20'W
Cajamarca
Celendín
Pass along the Cajamarca-Celendín rd.
Laguna Tembladera, Jalea de
3300
06°l4'S
79°l9'W
Lambayaque
Ferreñafe
Distrito Incahuasi, along Laguna Tembladera-Cerro Negro rd.
Laguna Yahuarcocha, Jalea de
3600
06° 15'S
79°l0'W
Cajamarca
Chota
Distrito Miracosta, above Incahuasi
Laguna Y anaco eha
4050
06°59'S
78°31'W
Cajamarca
Cajamarca
40 km N of Cajamarca, along rd. to Hualgayoc
Laguna de Culluna
3850
La Libertad
Pataz
Laguna de Santa Cruz Chiquita
4150
ors9'S
78°17'W
La Libertad
Santiago de Chuco
Near Shoreyo
Lagunas (Las)
4500
06°55'S
78°35'W
Cajamarca
Cajamarca
Ca. Km 50 on Cajamarca-Hualgayoc rd.
Lagunas Huarinjas, Pmo.
3800
05°02'S
79°29'W
Piura
Ayabaca
NWofSalalá
Lagunas Huaylillas
4000
orss·s
78°02'W
La Libertad
Huamachuco
N N N
See Jalea de Kumulea
Kumulka, Jalea de
2:':: tri
2:'::
o
~
(/l
o'Tl >-3
::r: tri
ztri
Lagunas de Las Compuertas, Jalea de
3750
Cajamarca
Cajamarca
Cajamarca--:Hualgayoc rd.
Laja, Pampa de la
3425
06°22'S
79°1l'W
Cajamarca
Chota
Between La Granja and Licupis
Llalladén, Cerro
4200
06°53'S
78°37'W
Cajamarca
Cajamarca
Ca. Km SI on Cajamarca-Hualgayoc rd.; possibly equals Las Lagunas
Llanacocha, Cerro
4225
or36'S
78°2l'W
Cajamarca
Cajabamba
o
Lluchubamba
4500
or3I'S
7r58'W
Cajamarca
Cajabamba
Lorito, Pampa de
3350
orr7'S
78°25'W
Cajamarca
Cajamarca
Luya Viejo
3300
06°08'S
7r59'W
Amazonas
Luya
Mahoma Chico, Cerro
3500
Cajamarca
Cajamarca
>-z ñ >1:-< o >-
Maqui Maqui, Jalea de
4120
06°57'S
78°28'W
Cajamarca
Cajamarca
Masma,La
3540
orws
78°14'W
Cajamarca
·cajamarca
E ofNamora
Michiquillay
3200
oro3'S
78°20'W
Cajamarca
Cajamarca
Encañada-Celendín rd.
Micuypampa
3600
oro2·s
78°14'W
Cajamarca
Migma
3600
or38°S
7r5r'W Cajamarca
Moneada
4900
or38'S
78°20'W
Morocha, La
3310
Cajamarca
Hualgayoc.
Motil, Jalea de
3500
La Libertad
Otuzco
Cajamarca
-< o
~
to
>-3
Above Huacraruco; also spelled Lirito
N of Cajamarca, along Shanta Alto rd.; includes a cerro and laguna by same name
Celendín
Km 62 on Cajamarca-Celendín rd.
Cajabamba
E ofCajabamba
Cajamarca
Minas (Las)
~
§ ztri
Cajamarca-Hualgayoc rd., before Las Lagunas Cajabamba Between Hualgayoc and Culquirumi
<o r
00
.¡;.
N ame Muchucaira, Jalea de
Elev. (m)
Latitud e
Longitude
4100
N anrá, Jalea de
Division 1
Division2
La Libertad
Otuzco
Cajamarca
Contumazá
Notes Heda. Motil-Santiago de Chuco, 1O km from Campero
Negritos
4050
06°54'S
78°32'W
Cajamarca
Cajamarca
N egro, Cerro (1)
3650
oroo·s
78°34'W
Cajamarca
Cajamarca
27 km a1ong the Cajamarca-Hualgayoc rd.
Negro, Cerro (2)
3650
05° IO'S
79°22'W
Cajamarca
San Ignacio
Near the border between Departamentos de Piura and Cajamarca
Ocumal
3370
06°27'S
78°06'W
Amazonas
Luya
Pajuela, La
3500
06°59'S
78°32'W
Cajamarca
Cajamarca
Km 30 along the Cajamarca-Hualgayoc rd.
Pampa Taguán
3500
06°56'S
78°13'W
Cajamarca
Celendín
Between Sorochuco and Agua Colorada
Paramillo, Cerro
3714
05°43'S
79°14'W
La Libertad
Paratón, Señal
3413
05°3l'S
79°28'W
Piura
'D 'D
::S
Km 35 along the Cajamarca-Hualgayoc rd.
Huancabamba See Jalea de J elig
Paso Gelig Paso de Alaska
See Alaska, Paso de 06°44'S
78°29'W
Cajamarca
3300
06°32'S
78°0l'W
. Amazonas
3220
oros·s
78°l7'W
Cajamarca
Cajamarca
Above Polloquito
3550
07°0l'S
78°38'W
Cajamarca
Cajamarca
Km 30 on Cajamarca-Hualgayoc rd., includes Porcón Alto
Piedras Gachas, Cerro
3900
Pisuquia Polloquito Porcón, Jaleas de
Cajamarca
Between Yanacocha and Llaucan
Luya
-o
See Jalea El Chuño
Pozo Chuño Pozo Kuán, Jalea del
4000
Cajamarca
Contumazá
Quesnada, Jalea de
4000
La Libertad
Santiago de Chuco
Quesquenda, Jalea de
4100
La Libertad
Santiago de Chuco
>~ 3:: o
[/)
Perhaps equals Jalea de Quesquenda
Quilimbach
3200
06°56'S
78°ll'W
Cajamarca
Celendín
Rd. to Celendín
Quilish
3500
oro2•s
78°34'W
Cajamarca
Cajamarca
N ofCajamarca, along rd. to Hualgayoc
Quinua, Cerro
3200
orm·s
78°13'W
Cajamarca
Celendín
Cajamarca--Celendín rd., toward Huagal
Quinuamayo
3520
oro3'S
78°l9'W
Cajamarca
Cajamarca
Between Encañada and Jalea de Kumulea
Quinuas, Jalea de Las
3900
or24'S
78°40'W
Cajamarca
Contumazá
Contumazá-Asunción rd.
Quiruvilca, Jalea de
4200
08°00'S
78° 19'W
La Libertad
Santiago de Chuco
Quesquenda, Lago El Toro; also spelled Quirubilca
Ramada, Jalea de La
3500
La Libertad
Huamachuco
20 km W ofHuamacucho
Rangra, Quebrada de
3600
La Libertad
Pataz
Tayabamba-Huangaspata
Rejo
3600
orOl'S
78°25'W
Cajamarca
Cajamarca
N ofRío Grande
Rodacocha
3700
07°0l'S
78°2l'W
Cajamarca
Cajamarca
Ruecas, Señal Cerro
4222
07"56'S
78°l5'W
La Libertad
Santiago de Chuco
N N
w
Elev. (m)
Latitud e
Longitude
Division 1
Division2
Rumi-Rumi
4496
or33'S
77°58'W
Cajamarca
Cajabamba
Rumiguacha, Cerro
4090
Cajamarca
Cajamarca
Sacsha
3850
07°01'S
78°29'W
Cajamarca
Cajamarca
Sal, Pampa de la
3500
or24'S
78°43'W
Cajamarca
Contumazá
Along Salcot-Cascabamba rd.
05°30'S
79°00'W
Cajamarca
Jaen
In range ofmountains NW ofJaen and SSE ofHuancabamba; coordinates are approximate
N ame
Sallique, Pmo. de
Notes
N N
.¡,..
Rocky jalea
San Antonio
3200
06°38'8
78°35'W
Cajamarca
Hualgayoc
13 km offthe Bambamarca-Chota rd.
San José, Jalea de Lagunas
3900
oroo·s
78°30'W
Cajamarca
Cajamarca
S ofLaguna Yanacocha
Sango, Cerro de
3900
La Libertad
Otuzco
Also spelled Cerro de Songo; Motil-Shorey rd., before Quiruvilca
Sansacocha,Laguna
3200
or47'S
7r59'W
La Libertad
Huamachuco
Sen dama!
3330
oroo·s
78° 12'W
Cajamarca
Celendín
Between Encañada and Celendín
[/)
Sexcemayo
3818
or12's
78°34'W
Cajamarca
Cajamarca
W ofCajamarca
...,
Shillas Negras, Cerro
3920
06°53'S
78°35'W
Cajamarca
Cajamarca
N ofLaguna Yanacocha
::r::
Shimbe, Cerro
3942
04 °58'S
79°27'W
Piura
Huancabamba
Near Laguna El Shimbe
ztT1
Shioglia
3700
06°57'S
78°36'W
Cajamarca
Cajamarca
Cajamarca-Hualgayoc rd.
Shoglla, Jalea de La
4000
Cajamarca
Hualgayoc
Sitacocha
3050
Soldaditos
3850
Suro
3800
or37'S
Tactama
3500
Tal langa, Jalea de Tambillos
Cajamarca
Cajabamba Cajamarca
High part of the Pajuela, turnoff at Km 30
7r56'W
Cajamarca
Cajabamba
Along rd. to Lluchubamba
78°14'W
06° 18'S
Amazonas
Luya
3600
or42'S
7r59'W
La Libertad
Sánchez Carrión
3850
or4o·s
78°20'W
Cajamarca
Cajabamba
Tambo
3650
oro9'S
78°07'W
Cajamarca
Cajamarca
Guanico-Guagal rd.
Tantahuatay, Cerro
3800
06°44'S
78°41'W
Turnoff along the Coymolache-Chugur rd.
Taulí, Jalea de
3900
Taulís Alto, Jalea
3100
Tingo, Cerro El
3350
Toro, Pampas del
06°44'S
7r58'W
78°40'W
Cajamarca
Hualgayoc
La Libertad
Pataz
Cajamarca
San Miguel
o
~
Cajamarca
orz9'S
;;:: tT1 ;;::
6 km from Marcaba! rd.; also called Hacienda Tallanga
o"rj
tT1
~
o"< ~ r: ttl o..., >z ñ >r' Cl
>-
Cajamarca
Hualgayoc
NW ofHualgayoc, toward Chugur
3400
Cajamarca
Celendín
Cajamarca-Celendín rd.
Totoraconga
3250
Cajamarca
Cajamarca
10 km NW ofCajamarca, along Chamis rd.
Totorilla
3650
or15'S
78°04'W
Cajamarca
San Marcos
NE ofSan Marcos
Travesía Laplap
3600
07°06'S
7r47'W
La Libertad
Bolívar
Between Longotea and Bolívar
§ tT1 z
<o r
00
.¡,..
N ame
Elev. (m) Latitude
Division 1 Cajamarca
Hualgayoc
Near Perlamayo
Cajamarca
Celendín
Between El Encañada and Celendín Along Balsas-Leimebamba rd.
Tres Lagunas
3300
Triunfo, Jalea El
3280
06°58'S
78°ll'W
Uchurama, Cerro
4317
06°51 'S
77°53'W
Ullauchan, Jalea de
3400
Usnio, Cerro El
3300
Vervena Victoria, La
Division2
Notes
Longitude
Amazonas
Chachapoyas
La Libertad
Otuzco
Chota-Shorey Distrito de la Encañada
07"03'S
78°19'W
Cajamarca
Cajamarca
3700
07"0l'S
78°26'W
Cajamarca
Cajamarca
4000
07"54'S
78°l7'W
La Libertad
Santiago de Chuco
Vilaya
3400
06°22'S
78°09'W
Amazonas
Luya
Viscachas, Cerro
4010
06°57'S
78°45'W
Cajamarca
Cajamarca
N ofCajamarca, vicinity Yanacocha; also spelled Vizcacha
Yamoca, Pmo. de
2350
Cajamarca
Bracamora
Near San Felipe
Yanaguanaga,Prno.de
3100
Yerbabuena
3400
Cajamarca 07"3!'S
78°20'W
Cajamarca
'-0 '-0
..::;
Rd. to Conzuzo, area around the lake
Bonpland collection at Paris Cajabamba
.,:>
S:
~
o
(/)
N
N V1
226
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
[VOL. 84
Bibliographic Sou.rces for Locations of Páramos General Hanson, E. P. & N. Raymond (eds.). 1945. Index to maps of Hispanic America. Maps of Hispanic America. 1:1,000,000. American Geographical Society, Publication No. 5. U.S. Govemment Printing Office, Washington, OC.
Costa Rica American Geographical Society ofNew York. 1937. L¡go de Nicaragua, Central America. 1:1,000,000. Map NC16. Provisional edition. A. Hoen & Co., Baltimore. National Geographical Society. 1939. Mexico, Central America and the West Indies. 1:5,702,000. Cartographic Division, Washington, OC. - - - . 1986. Central America. 1:2,534,000. Cartographic Division, Washington, OC. Rand McNally & Co. 1969. Costa Rica. Mapa vial. Lithographed in U.S.A. Tosi, J. A., Jr. 1969. República de Costa Rica, mapa ecológico. Centro Científico Tropical. 1:750,000. Litografiado por Instituto Geográfico Nacional. San José, Costa Rica. Weber, H. 1959. Los páramos de Costa Rica y su concatenación fitogeográfica con los Andes suramericanos. Instituto Geográfico de Costa Rica, San José. - - - . 1963. Über die Vegetation der hochandinen Páramos. Pp. 2-16. Jahrbuch 1963 des Vereins zum Schutze der Alpenpflanzen und Tiere e. V. (Bd. 28), Munich.
Pan ama American Geographical Society of New York. 1928. Panama, Central America. 1:1,000,000. Map NC-1 7. Provisional edition. A. Hoen & Co., Baltimore. Instituto Costaricense de Turismo. 1977. Panamá. Mapa vial. Litografia Trejos, Costa Rica. Instituto Geográfico Nacional "Tommy Guardia." Panamá. Plano de la ciudad de Panamá y mapas de carreteras. National Geographical Society. 1939. Mexico, Central America and the West Indies. 1:5,702,000. Cartograpic Division, Washington, DC. Rand McNally & Co. 1967. Panamá. Mapa vial. Lithographed in U.S.A.
Colombia American Geographical Society ofNew York. 1942. Cali. Map NA-18. 1:1,000,000. Provisional edition. A. Hoen & Co., Baltimore. - - - . 1944. South America. Series map ofthe Americas, Sheet lB. Second edition (copyright 1942). A. Hoen & Co., Baltimore. - - - . 1945. Bogotá. Map NB-18. 1:1,000,000. Provisional edition. A. Hoen & Co., Baltimore. - - - . 1952. Barranquilla. Map NC-18. 1:1,000,000. Provisional edition. A. Hoen & Co., Baltimore.
Anonymous. 1:100,000 topographic maps Colombia [Map No. 98, Durania; 110, Pamplona; 121, Cerrito; 137, El Cocuy; 153, Chita; 172, Paz del Río; 192, Laguna de Tota; 228, Bogotá d.e. Noreste; 280, Palmira; 300, Cali; 364, Timbío; 387, Bolívar; 429, Pasto]. Anonymous. Colombia maps [Map No. 2-060, Tunja]. Becking, M. L., A. J. Negret & R. G. M. Hofstede. 1997. Caracterización preliminar de un paramillo en la Cordillera Occidental del Departamento del Cauca. Noved. Colomb. (Mus. Hist. Nat. Univ. Cauca) 7: 33-44. Cleef, A. 1980. La vegetación del páramo neotropical y sus lazos Australo-Antárticos. Colombia Geogr. 7(2): 7-49. Corporación Autónoma Regional del Cauca - Oficina de Planeación. 1983. Recreación en la área del Alto Cauca. Recursos espaciales en el área rural. Elaboración, diseño, dibujo y diagramación por Unidad de Desarrollo Regional y Urbano, Oficina de Planeación, CVC, Cali. Cuatrecasas, J. 1969. Prima flora colombiana. 3. CompositaeAstereae. Webbia 24: 1-335. - - - . 1985. Brunelliaceae. Fl. Neotr. Mono gr. 2(Suppl.): 1-103. - - . 1989. Relación de las localidades y fechas correspondientes a las plantas colectadas y anotadas por José Cuatrecasas, al servicio del Instituto de Ciencais Naturales (Herbario Nacional Colombiano- COL) (V-1939IX-1942). Perez-Arbelaezia 2: 293-328. Epling, C. & Játiva, C. 1964. Revisión del género Satureja en América del Sur. Brittonia 16: 393-416. Hettner, A. 1892. Die Kordillere von Bogotá. Petermanns geographische Mitteilungen, Supp. 104 [Original pagaination unknown]. [Translated into Spanish by E. Guhl as: La Cordillera de Bogotá: Resultados de viajes y estudios. 351 pp. 1966. Banco de la República, Bogotá.] INDERENA. 1986. Puracé: Montaña de fuego, nieve, páramo y selva. Revista Parq. Nac. Colombia 1(9):1-16. Instituto Geográfico "Agustín Codazzi." 1963. Carta preliminar (blueprint) maps. 1:25,000 [Plancha 152-IIIB, C, andO]. - - - . 1968. República de Colombia." Mapa físicopolítico. 1:1,500,000. Edición preliminar. Instituto Geográfico "Agustín Codazzi," Bogotá. - - - . 1971. Diccionario geográfico de Colombia. Tomo I (Letras A-L) y II (Letras M-Z). Ministerio de Hacienda y Crédito Público. Edición patrocinada por el Banco de la República, Bogotá - - - . 1972. Carta general. Plancha N° 9. Carta preliminar. 1:500,000. Ministerio de Hacienda y Crédito Público, Bogotá. - - - . 1973. Mapa vial. Basado en el mapa de la República de Colombia a escala 1:1 ,500,000 del Instituto Geográfico "Agustín Codazzi." Ministerio de Hacienda y Crédito Público. Carvajal & Cia., Bogotá. - - - . 1976. Colombia departamentos, intendencias y comisarías: Información geográfica básica. Bogotá. - - - . 1977. Atlas de Colombia. Tercera edición. Litografia Arco, Bogotá.
1999]
PARAMOS
- - - . 1983. República de Colombia. Mapa fisico-político. 1:1,500,000. Edición duodécima. Instituto Geográfico "Agustín Codazzi," Bogotá. - - - . 1985. Mapa de la República de Colombia. Séptima edición. Carvajal S.A., Colombia. Killip, E. P. 1927. Report ofthe Killip-Smith botanical expedition to Colombia, 1926-27. J. New York Bot. Gard. 28: 205-220. Llano, M. del. 1990. Los páramos de los Andes. Montoya & Araújo Ltda., Bogotá. Mathias, M. E. & Constance, L. 1962. The Andean genus Niphogeton (Umbelliferae)revisited. Brittonia 14: 148-155. Meyer de Schauensee, R. 1948-1951. The birds ofthe Republic ofColombia. Pts. 1-4. Caldasia 5(22-26): 251-1112. Paynter, A. R., Jr. & Traylor, M. A. 1981. Omithological gazetteer of Colombia. Bird Department, Museum of Comparative Zoology, Harvard University, Cambridge. Reyes Z., P., J. Molano B., F. G. A. Cortés Lombana, J. O. Rangel Ch., A. Flórez, P. Iriarte & E. Kraus (eds.). 1995. El páramo: Un ecosistema de alta montaña. Serie Montañas Tropoandinas, Vol. l. Fundación Ecosistemas Andinos (ECOAN), Santa Fé de Bogotá. Robson, N. K. R. 1987. Studies in the genus Hypericum L. (Guttiferae) 7. Section 29. Brathys (Part 1). Bull. Brit. Mus. Nat. Hist. (Bot.) 16: 1-106. Sánchez P., H. & J. Hernández-Camacho. 1995. Labiodiversidad de los Andes de Colombia y su conservación en los parques nacionales. Pp. 619-626. In: S. P. Churchill, H. Balslev, E. Forero & J. L. Luteyn (eds.), Biodiversity and conservation of neotropical montane forests. The New York Botanical Garden, Bronx. Smith, A. C. & M. F. Koch. 1935. The genus Espeletia: A study in phylogenetic taxonomy. Brittonia 1: 479-530. Sturm, H. & O. Rangel Ch. (eds.). 1985. Ecología de los páramos andinos: Una visión preliminar integrada. Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bibliot. José Jerónimo Triana 9: 1-292. United States Board on Geographical Names. 1988. Gazetteer of Colombia. Third edition. Defense Mapping Agency, Washington, DC. Viloria P., A. L. 1993. Los páramos de Perija. Natura 93: 25-29. Revista de Divulgación Científica Sociedad de Ciencias Naturales La Salle, Caracas. Wood, J. R. I. & Harley, R. M. 1988. The genus Salvia (Labiatae) in Colombia. Kew Bull. 44: 211-278.
Venezuela American Geographical Society ofNew York. 1945. Map CN-19, Caracas. Compiled and drawn by The American Geograpichal Society ofNew York. Aranguren B., A. & N. J. Márquez. 1995. Use, collection, commercialization, and vulnerability of two species of the genus Oritrophium (0. venezuelense and O. peruvianum) in the Venezuelan Andes. Final Report. Programa Andes Tropicales, Fundación BIOMA, Mérida, Venezuela. - - - , - - - , R . Prato & Y. Lesenfants. 1996. Use, collection, commercialization, and vulnerability oftwo species of the genus Oritrophiwn (O. venezuelense and O. peruvianum, Compositae) in the Venezuelan Andes. Acta Bot. Venez. 19: 16-38. Bono, G. 1996. Flora y vegetación del Estado Táchira, Venezuela. Monografie 20. Museo Regionale di Scienze Natrurali, Torino.
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Caribbean Petroleum Company. 1943. Mapas de las vías de comunicación de los Estados Unidos de Venezuela. 1:2,000,000. Proyección plicónica. Top. Dept. C.P.C., Maracaibo. Gabaldón, M. 1992. Parques nacionales de Venezuela. Serie Parques Nacionales y Conservación Ambiental, Caracas. Jahn, A. 1912. La cordillera venezolana de los Andes. Tipografía Comercio, Caracas. (Cited in Pérez, 1992a.] - - - . 1931. Los páramos venezolanos: Sus aspectos fisicos y su vegetación. Bol. Soc. Venez. Ci. Nat. 1(3): 93127. López del Pozo, E. 1993. El páramo: Diferentes visiones. Pp. 109-121. In: C. Schubert & L. Vivas (eds.), El Cuaternario de la Cordillera de Mérida (Andes venezolanos). Universidad de Los Andes/Fundación Polar, Mérida. López Figueiras, M. 1986. Censo de macrolíquenes venezolanos de los estados Falcón, Lani, Mérida, Táchira y Trujillo. Universidad de Los Andes, Mérida. Ministerio de Obras Públicas. 1969. Atlas de Venezuela. Dirección de Cartografia Nacional, Caracas. - - - . 1969-1977. Maps of Venezuela. Serie DCN. 1:25,000. Dirección de Cartografia Nacional, Caracas. [Hoja 5839, San José de Bolívar. 1976; Hoja 5840, La Grita. 1976; Hoja 5841, El Vigía. 1977; Hoja 5940, Libertad. 1976; Hoja 5941, Mérida. 1977; Hoja 5942, La Azulita. 1977; Hoja 6041, Ciudad de Bolivia. 1977; Hoja 6042, Timoles. 1976; Hoja 6142, Barinitas. 1975; Hoja 6143, Boconó. 1977; Hoja 6144, Trujillo. 1969; Hoja 6244, Biscucuy. 1976; Hoja 6245, El Tocuyo. 1975; Hoja 6043, Valera. 1976.] - - - . 1977. Mapa de las carreteras de Venezuela. 1:1,000,000. Octava edición. Dirección General de Vialidad. Gráficas Artimano C.A., [Caracas] Venezuela. - - - . [Without date.] Mapa de las carreteras de Venezuela, Región Norte de la República de Venezuela. 1:1,000,000. Dirección General de Vialidad. Gráficas Artimano C.A., [Caracas] Venezuela. Ministerio del Ambiente y de Los Recursos Naturales Renovables. 1978. Mérida. Map NC 19-13. 1:250,000. Dirección General de Información e Investigación del Ambiente, Dirección Nacional de Cartografia- MARNR. Caracas. - - - . 1983. Venezuela en mapas. Primera edición. Dirección General de Información e Investigación del Ambiente, Dirección de Cartografia- MARNR. Caracas. Monasterio, M. (ed.). 1980. Estudios ecológicos en los páramos andinos. Ediciones de la Universidad de Los Andes, Mérida, Venezuela. Paynter, A. R., Jr. 1982. Ornithological gazetteer o[ Venezuela. Bird Department, Museum ofComparative Zoology, Harvard University, Cambridge. Schubert, C. 1976. Evidence of former glaciation in the Sierra de Perijá, western Venezuela. Erdkunde 30: 222--224. Sívoli G., A. [Date uncertain but 1976 or earlier]. Diccionario geográfico de Venezuela. Ediciones Eneva, C.A., Caracas. Unitcd States Board on Geographical Names. 1961. Venezuela. Gazetteer No. 56. Office of Geography, Department ofthe Interior, Washington, DC. Vila, M. A. 1976. Diccionario de tierras y aguas de Venezuela. Ministerio de Obras Públicas, Dirección de Cartografia Nacional, Caracas. Viloria P., A. L. 1993. Los páramos de Perijá. Natura 93: 25-29. Revista de Divulgación Científica Sociedad de Ciencias Naturales La Salle, Caracas.
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MEMOIRS OF THE NEW YORK BOTANICAL GARDEN
Vuilleumier, F. & D. N. Ewert. 1978. The distribution of birds in Venezuelan páramos. Bull. American Mus. Nat. Hist. 162(2): 47-90. Xena de Enrech, N. 1992. Valerianaceae. Fl. Venezuela 5(1 ): 221-267.
Ecuador Acosta-Solís, M. 1980. Plantas indígenas para forrajicultura tropandina.' Revista Acad. Colomb. Ci. Exact. 15(56): 57-97. - - - . 1984. Los páramos andinos del Ecuador. Publicaciones Científicas MAS. Quito. American Geographical Society ofNew York. 1938. Quito. Map SA-17. 1:1,000,000. Provisional edition. A. Hoen & Co., Baltimore. - - - . 1938. Piura. Map SB-17. 1:1,000,000. Provisional edition. A. Hoen & Co., Baltimore. - - - . 1949. Río Mira-Islas Galápagos. Map NA-18, Southamerica. 1:1,000,000. Provisional edition. A. Hoen & Co., Baltimore. Andrade Marín, L. 1952. La desconocida región de Oyacachi: Rectificaciones geográficas, hallazgos etnológicos y de un precioso manuscrito inédito, en poder oculto de los indios. Anales Organo Univ. Central (Ecuador) 79(331-332): 5-63. Diels, L. 1937. Beitrage zur Kenntnis der Vegetation und Flora von Ecuador. Biblioth. Bot. 116: 1-190. [Spanish translation as: Contribuciones al conocimiento de la vegetación y flora del Ecuador. Universidad Central, Quito. August, 1938.] Instituto Geográfico Militar. 1950. Mapa geográfico del Ecuador. 1:1,000,000. Quito. - - - . 1978. Indice toponímico de La República del Ecuador. Tomo I-VII. Quito.
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Mena V., P. 1995. Las áreas protegidas con bosque montano en el Ecuador. Pp. 627-635. In: S. P. Churchill, H. Balslev, E. Forero & J. L. Luteyn (eds.), Biodiversity and conservation of neo tropical montan e forests. The New York Botanical Garden, Bronx. Paynter, A. R., Jr. & M. A. Traylor. 1977. Ornithological gazetteer of Ecuador. Bird Department, Museum of Comparative Zoology, Harvard University, Cambridge. Sampedro F., Ing. Capt. 1950. Mapa del Ecuador. 1:1,000,000. United States Coast and Geodetic Survey. - - - . 1976. Atlas geográfico del Ecuador. Artes Gráficas, Quito. United States Board on Geographical Names. 1987. Gazetter of Ecuador. Ed. 2. Defense Mapping Agency, Washington, De.
Peru American Geographical Society of New York. 1949. !quitos. Map, SA-18. Provisional edition. A. Hoen & Co., Baltimore. - - - . 1938. Piura. Map SB-17. 1:1,000,000. Provisional edition. A. Hoen & Co., Baltimore. CartaNacional. 1:100,000. Dillon, M. O. & A. Sagástegui A. 1986. Jalcophila, a new genus of an andean Inuleae (Asteraceae). Brittonia 38: 162-167. Herrera de Laja, B. 1980. Revisión de las especies peruanas del género Gynoxys. Bol. Soc. Peruana Bot. 8(1, 2): 3-74. Instituto Geográfico Nacional. Mapas fisico-político. [Piura 1:500,000; Lambayeque 1 :3,000,000; Cajamarca 1:480,000; La Libertad 1:420,000; Amazonas 1:600,000.] Smith, D. N. 1988. Flora and vegetation ofthe Huascarán National Park, Ancash, Peru, with preliminary taxonomic studies for a manual ofthe flora. Ph.D. dissertation. Iowa State University, Ames.
BOTANICAL LITERATURE REFERENCES TO PARAMO
This bibliography of páramo literature is primarily botanical in orientation. Also included are a few studies ofpuna or afro-alpine (and other Old World) regions for comparati ve purposes, anda few zoologically oriented páramo studies. Although literature searches for these latter subjects have not been exhaustive, I have tried to add relevant papers with extensive bibliographies whenever possible. I have also made an effort to include páramo-based unpublished theses for which I have looked diligently in the various libraries ofthe Universidad de Los Andes (Facultad de Ciencias, Mérida, Venezuela), Universidad Nacional de Colombia (Instituto de Ciencias Naturales, Bogotá, and Facultad de Ciencias Exactas y Naturales, Medellín), Pontificia Universidad Católica del Ecuador (Departamento de Biología, Herbario QCA, Quito), and the University of Amsterdam (Rugo de Vries Laboratory, The Netherlands). I have not included in this bibliography strictly taxonomic revisions or monographs in which páramo species are a major element; they are, instead, included earlier in this book, in the "Checklist of Páramo Plants," and are cited with their respective families or genera. Although neither algae nor fungi (with the exception of lichenized fungi) are included in the abo ve checklist, literature references to them are included below. As
an additional help, at the end of most bibliographic entries, after the long dash (-), I have tried to note the main emphasis of the cited work, using the following list of subject categories.
Aagaard, E. M. J. 1982. Ecological distribution of mammals in the cloud forests and páramos of the Andes, Mérida, Venezuela. Ph.D. dissertation. Colorado State University. [Cited in J. F. Eisenberg, Mammals of the Neotropics: The northern Neotropics. VoL l. University ofChicago Press, Chicago.]-1, 4 Abadia, S. 1943. Cultivo de pastos de tierra fria. Bol. Ganadería 5: 333-373. (Cited in Roseveare, 1948.]-1, 10 Acosta de Obando, R. M. 1982. Estudio anátomo-foliar del género Carramboa Cuatr. Revista Fac. Farmacia (Universidad de Los Andes, Mérida, Venezuela) 22: 91129.-11 Acosta Saignes, M. 1952. El área cultural prehispánica de Jos Andes venezolanos. Archivos Venezolanos de Folklore 1(1): 45-80. [Cited in López del Pozo, 1993.]-5, JO, 14 Acosta-Solís, M. 1937. Excursión botánica al Páramo del Angel. Flora (Quito) 1(2): 103-118.-1, 5 - - - . 1942. El Quishuar u olivo del páramo: Olivo de Jos Incas. Flora (Quito) 2(5-6): 119-125.-1,2, 10
- - - . 1957. Clasificación geobotánica de los bosques y las otras formaciones vegetales del Ecuador. Ci. & Nat. 1(2): 62-77.-1 - - - . 1960. Los pastizales naturales del Ecuador: Conservación y aprovechamiento de los páramos y sabanas. Revista Geogr. (Rio de Janeiro) 53: 87-99. -1 - - - . 1962a. Terminología geográfica y ecológica para América Tropical Andina. Revista Acad. Colomb. Ci. Exact. 11(44): 351-358.-1, 5 - - - . 1962b. Fitogeografia y vegetación de la Provincia de Pichincha. Publicación 249: 1-13 5. Instituto Panamericano de Geografia e Historia, México, D.F. -1,9 - - - . 1965. Los recursos naturales del Ecuador y su conservación. Parte 1' (Pub l. 254) y 2' (Tomo I) (Pub l. 255). Instituto Panamericano de Geografia e Historia, México, D.F. -1, 2, 5, 6 - - - . 1966. Las divisiones fitogeográficas y las formaciones geobotánicas del Ecuador. Revista Acad. Colomb. Ci. Exact. 12(48): 401-447. -1, 5, 9
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2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19
ecology (including geoecology, reproductive biology, community ecology, phyto-sociology, population and vegetation dynamics, etc.) systematics physiology, ecophysiology zoology general (including natural history, travels, etc.) conservation geology (including soiis, geologic history, glaciation, etc.) palynology, paleoecology, and paleoclimatology biogeography ethnobiology (including agricultura! systems, land use and management) anatomy & morphology chemistry & pharmacology archeology anthropology (including sociology, human ecology) limnology present-day climatology (including microclimatology) algae fungi (including lichens) bryophytes
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México, y su posición en las montañas tropicales de ---·. 1968a. Protección y conservación de la naturaleza en Sudamérica. Vol. 1: 230-250. In: Fittkau et al. (eds.), América. Phytocoenologia 22: 391-436.-1, 5, 9 1968.-6 Alulima Gordillo, J. R. & J. Roberto. 1993. Formas ade- - - . 1968b. Divisiones fitográficas y formaciones geocuadas de propagación vegetativa y repoblación arborea de Alnusjorullensis, Buddleia in cana y Polylepis in cana botánicas del Ecuador. Casa de la Cultura Ecuatoriana, Quito. -1, 5, 9 para las comunidades rurales del Chimborazo. Tesis de - - - . 1969. Glumifloras del Ecuador: Catálogo fitogeoIngeniero Forestal. Universidad Nacional de Loja, Loja. -1,6,14 gráfico de las Gramíneas, Ciperáceas y Juncáceas. Flora Al varado, L. 1945. Glosario del Bajo Español en Venezuela. (Quito) 13(47-50): l-216. -1,2 Primera parte-Acepciones especiales. Ministerio de - - - . 1973. El paisaje y la cubierta vegetal del Reino de Educación, Dirección de Cultura y Bellas Artes, Caracas. Quito al arribo de los conquistadores españoles. Bol. Inform. Ci. Ecuador 14(105-106): 1-17.-5, lO, 14 [Cited in López del Pozo, 1993.]-5, 14 - - - . 1977. Ecología y fitoecología. Casa de la Cultura Amat García, G. 1987. Influencia del uso del suelo sobre la Ecuatoriana, Quito. -1 mesofauna edáfica en el Páramo de Monserrate, - - - . 1980. Plantas indígenas para forrajicultura tropanCundinamarca. Tesis. Dpto. de Biología, Universidad Nacional de Colombia, Bogotá. -1, 4 dina. Revista Acad. Colomb. Ci. Exact. 15(56): 57-97. -10 ----. 1991 a. Artropofauna del Parque Nacional Natural - - - . 1982. Los pastizales naturales del Ecuador: Conser. · Chingaza. Cuad. Divulg. 14: 1-13.--4 vación y aprovechamiento de los páramos y sabanas. ·-·- - . 1991 b. Caracterización de microhábitats de la artroRevista Geogr. (Quito) 17:87-99.-1,6 pofauna en páramos del Parque Nacional Natural - - - . 1984. Los páramos andinos del Ecuador. Pub!. Chingaza, Cundinamarca, Colombia. Caldasia 16(79): Cient. MAS, Quito. -1, 2, 5, 6 539-550. (Contains good bibliography.]-1, 2, 4 - - - . 1985. El arenal del Chimborazo, ejemplo de puna Andrade, C., N. Aranguren, G. Cardenas, H. Florindo, W. en el Ecuador. Revista Geogr. (Quito) 22: 115-122.-1 López, G. Oquendo, P. Patiño & G. Rueda. 1992. Estudio - - - . 1994. Los pajonales de Chiquicagua y la protección limnológico de tres lagunas en el Páramo de Chisacá del Río Colorado. RevistaGeogr. (Quito) 33: 7-18.-1,6 (Cundinamarca). Tésis de grado. Universidad Pedagó- - - & S. T. Machado. l937a. Estudio botánico-químico gica Nacional, Bogotá, Colombia. [Cited in Gonz.ález del Espeletia grandijlora (frailejón del Páramo del AnGonzález, 1995.]-15 gel). Instituto Botánico de la Universidad Central, Quito, Andrade, G. I. (ed.). 1993. Carpanta: Selva nublada y Ecuador. -1, 2, 5, 10, 12 páramo. Fundación Natura Colombia, Bogotá. -5 - - - & - - - . 1937b. Espeletia grandijlora (E. hartAndrade Marín, L. 1936. Viaje a las misteriosas montañas wegiana), "Frailejón" del Páramo del Angel. Anales de Llanganati. Ecuadorian Institute ofNatural Sciences, Univ. Centr. Ecuador 59(301): 165-226.-1, 5 Quito. (Cited in Harling, 1979.]-5 Adams, M. J. 1973. Ecological zonation and the butterflies - - - . 1952. La desconocida región de Oyacachi: Rectiofthe Sierra Nevada de Santa Marta, Colombia. J. Nat. ficaciones geográficas, hallazgos etnológicos y de un Hist. 7:699-718.-1,4 precioso manuscrito inédito, en poder oculto de los Aguado, Fray Pedro de. 1581. Recopilación historial de indios. Anales Organo Univ. Central (Ecuador) 79(331Venezuela. Reprinted in 1963 as part of the series 332): 5-63.-5,14 Fuentes para la Historia Colonial de Venezuela, vols. Andressen, R. & Ponte, R. 1973. Estudio integral de las 62 and 63. Biblioteca de la Academia Nacional de la Hiscuencas de los ríos Chama y Capazón: Climatología e toria, Caracas. [Cited in López del Pozo, 1993.]-5, 14 hidrología. Sub-proyecto No. II, Instituto de Geografía Aguirre C., J., E. Linares, E. Santana & G. Castro. 1986 y Conservación de Recursos Naturales, Universidad de [Abstract]. La brioflora de los alrededores de Bogotá. P. Los Andes, Mérida. [Cited in Pérez, 1995b.]-16 194. Resúmenes IV Congr. Latinoamer. Bot. (Medellín, Andriessen, P. A. M., K. F. Helmens, H. Hooghiemstra, P. Colombia). -1, 2, 19 A. Riezebos & T. Van der Hammen. 1993. Absolute - - - & O. Rangel Ch. 1976. Contribución al estudio ecochronology of the Pliocene-Quatemary sediment selógico y fitosociológico de las comunidades acuáticas quence ofthe Bogotá area, Colombia. Quatem. Sci. Rev. macroscópicas y continentales del Lago de Tota (Boyacá) 12: 483-501. [Also published in: Hooghiemstra (ed.), y alrededores. Tesis de grado. Dpto. de Biología, UniThe Quaternary ofColombia, 22. 1996.]-8 versidad Nacional de Colombia, Bogotá. -1, 3, 15 Anés, J. & O. Crescente. 1997 [Abstrae!]. 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1999]
PARAMOS
Yánez M., A. P. 1997. Análisis de la distribución de especies vegetales a lo largo de un gradiente altitudinal páramoselva nublada del Parque Nacional Sierra Nevada, Venezuela. Tesis de M.Sc. Facultad de Ciencias, Universidad de Los Andes, Mérida. -1 Yerena, E. 1994. Corredores ecológicos en los Andes de Venezuela. Parques Nacionales y Conservación Ambiental No. 4. Fundación Polar and Instituto Nacional de Parques (INPARQUES), Caracas. -1, 4, 5, 6, 9 ~~-. 1997. Corredores ecológicos en los Andes de Venezuela. Revista Ci. UNET (Universidad Nacional Experimental del Táchira) 9(2): 42-45. -6, 9 Young, K. R. 1993a. Woody and scandent plants on the edges of an Andean timberline. Bu! l. Torrey Bot. Club 120: 1-18.-1 ~~-. 1993b. National park protection in relation to the ecological zonation of a neighboring human community: An example from northern Peru. Mountain Res. Developm. 13: 267-280.-1, 10 ~~-. 1993c. Tropical timberlines: Changes in forest structure and regeneration between two Peruvian timberline margins. Arctic Alpine Res. 25: 167-174.-1, 10 ~~-. 1994. Roads and the environmental degradation of tropical montane forests. Conservation Biol. 8: 972-976. -1, 6,10 ~~-. 1997. Wildlife conservation in the cultural landscapes ofthe central Andes. Landscape and Urban Planning 38: 137-147.-5, 6, 10, 14 ~~- & B. León. 1988. Vegetación de la zona alta del Parque Nacional Río Abiseo, San Martín. Revista Forestal Perú 15: 3-20. -1 ~~- & ~~-. 1990. Catálogo de las plantas de la zona alta del Parque Nacional Río Abiseo, Perú. Pub!. Mus. Hist. Nat. Univ. Nac. Mayor San Marcos, ser. B, 34: 1-37.-2
~~-
271
& ~~-. 1991. Diversity, ecology & distribution of high-elevation pteridophytes within Río Abiseo National Park, north-central Peru. Fem Gaz. 14: 25-39.-1,2 ~~- & C. Reynel. 1997. Huancabamba region, Peru and Ecuador. Pp. 465-469. In: Davis et al. (eds.), 1997.1, 2, 5, 6, 9 (regional account) ~~-,B. León, A. Cano & O. Herrera-MacBryde. 1997. Peruvian puna, Peru. Pp. 470-476. In: Davis et al. (eds.), 1997. -1, 2, 5, 6, 9 (regional account) Zadroga, F. 1972. Sorne observations on man's impact on sorne natural and man-made bogs in the area of El Empalme, Talamanca range, Costa Rica. Pp. 125-146, in the coursebook for the geography field course. Organization for Tropical Studies, San José, Costa Rica. [Cited in Hom, 1988.]-14 Zambrano, C. V. (ed.). 1993. Hombres de páramo y montaña: Los yanaconas del Macizo Colombiano. Instituto Colombiano de Antropología, COLCUL TURA y PNR, Bogotá. -5, 10,14 Zerries, O. 1968. The South American Indians and their culture. Pp. 329-388. In: Fittkau et al. (eds.), 1968. Zimmerer, K. S. & K. R. Young (eds.). In press. Nature's geography: New lessons for conservation in developing countries. University of Wisconsin Press, Madison. -1,6, 10,14 Ziittle, H. W. 1970. Die Eisendynamik in Biiden der PáramoStufe der Anden Venezuelas. Zeitschr. Pflanzenernahrung Biidenkunde 127: 10-18.-1,7 Zuluaga, J. E. & L. Mattson. 1981. Glaciaciones en la Cordillera Occidental de Colombia, Páramo de Frontino, Departamento de Antioquia. Mem. I Sem. Cuat. Colombia, Revista CIAF (Bogotá) 6(1/3): 400-440, 639-654. [Cited in García Velez & Londoño, 1985; and also in Kroonenberg et al., 1990.]
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Appendix 1: Alphabetical List ofthe Families and Genera in the Pรกramo Checklist Numbers in parentheses after family names signify the number of genera/species represented in pรกramo for that family. The number in parentheses after generic names signifies the number of species represented in pรกramo for that genus. Lichens Lichenized Ascomycetes Acarosporaceae ( 1/1) Polysporina (1) Arthophyreniaceae (1/1)
Mycomicrothelia ( 1) Arthrorhaphidaceae (1/2) Arthrorhaphis (2) Asterothyriaceae (1/1)
Asterothyrium (1) Bacidiaceae (4/9) Bacidiospora (1) Hypocenomyce (1) Phyllospora (4) Tephromela (3) Baeomycetaceae (2/4) Baeomyces (1) Phyllobaeis (3) Caliciaceae (113) Calicium (3) Cande1ariaceae (2/2) Candelaria ( 1) Candelariella ( !) Catillariaceae (2/2) Lopezaria (1) Sporastatia (1) Chrysothrichaceae ( 111) Chrysothrix (1) Cladoniaceae (3/45) Cladia (2) Cladina (5) Cladonia (38) Coccocarpiaceae (1/2) Coccocarpia (2) Collemataceae (2/26) Collema (1) Leptogium (25) Coniocybaceae (l/7) Chaenotheca (7) Ectolechiaceae ( 112) Tapellaria (2) Fuscideaceae ( 111) Maronea (1) Gya1ectaceae ( 111) Dimerella (1) Gomphillaceae (3/4) Echinoplaca (!) Gomphillus ( 1) Gyalideopsis (2) Hymene1iaceae (2/2) Megaspora (!) Tremolecia ( 1)
Icmadophi1aceae (2/4) Dibaeis (3) Icmadophila (1) Lecanora~eae (4/J]) Carbonea ( 1} Lecanora (3) Lecidella (6) Rhizoplaca ( 1) Lecideaceae (115) Lecidea (5) Lobariaceae (3/24) Lobaria (6) Pseudocyphellaria (5) Sticta (13) Mega1osporaceae (113) Megalospora (3) Pannariaceae (4/1 O) Erioderma (4) Pannaria (2) Parme/iella (2) Psoroma (2) Parmeliaceae (251159) Alectoria (2) Anzia (5) Bulbothrix (2) Canoparmelia (!) Cetraria (4) Cetrariatrum (3) Cetrariella (!) Everniastrum (1 O) Everniopsis (1) Flavoparmelia (3) Flavopunctelia ( 1) Hypogymnia (2) Hypotrachyna (50) Melanelia (1) Oropogon (21) Parmelina (1) Parmelinopsis (3) Parmotrema (13) Protoparmelia (2) Psiloparmelia ( 1) Punctelia (3) Rimclia (2)
Tuckneraria (1) Usnea (8) Xanthoparmelia (18) Peltigeraceae (3/ 13) Nephroma (1) Peltigera (1 O) So/orina (2) Pertusariaceae ( 1/2) Pertusaria (2)
Ph1yctidaceae ( 111) Phlyctis (1) Physciaceae (8/27) Buellia (1) Dirinaria (1) Hafellia (1) Heterodermia (17) Hyperphyscia (1) Phaeophyscia (2) Physcia (3) Pyxine (1) Pilocarpaceae ( 1/1) Byssoloma (1) Porpidiaceae (1/1) Porpidia (1) Psoraceae ( 1/1) Psora (1) Ramalinaceae (1/12) Ramalina (12) Rhizocarpaceae ( 1/2) Rhizocarpon (2) Solorinellaceae (1/2) Gyalidea (2) Sphaerophoraceae (1/1) Bunodophoron (1) Stereocau1aceae (l/19) Stereocaulon (19) Teloschistaceae (3/7) Ca/aplaca (2) Teloschistes (4) Xanthoria (1) Thelotremataceae (2/5) Diploschistes (4) Thelotrema (l) Trapeliaceae (4/8) Placopsis (2) Placynthiella (1) Trapelia (3) Trapeliopsis (2) Umbilicariaceae (1/9) Umbilicaria (9) Verrucariaceae (2/4) Catapyrenium (3) Normandina (1) Incertae Sedis Cystocoleus (l)
Lepraria ( 1) Leprocaulon (4) Lepra/ama (l) Racodium (1) Siphula (3) Thamnolia (l)
1999] Lichenized Basidiomycetes Meruliaceae (1/2) Dictyonema (2) Tricholomataceae (2/4) Omphalina (2) Phytoconis (2)
Mosses Ade1otheciaceae (111) Adelothecium ( 1) Amblystegiaceae (9119) Calliergon (4) Calliergone/la ( 1) Campyliadelphus (!) Cratoneuron (1) Drepanocladus (6) Gradsteinia (1) Hygroamblystegium (1) Sanionia ( 1) Scorpidium (3) Andreaeaceae (211 O) Acroschisma ( 1) Andreaea (9) Aulacomniaceae (111) Au/acomnium (1) Bartramiaceae (7/40) Anacolia (1) Bartramia (8) Breutelia (13) Conostomum ( 1) Flowersia ( 1) Leiome/a ( 6) Philonotis (1 O) Brachytheciaceae (7118) Aerolindigi& (l) Brachythecium (9) Eurhynchium (1) Pa/amocladium (l) Platyhypnidium (!) Rhynchostegium (3) Rozea (2) Bruchiaceae ( 111) Eobruchia (1) Bryaceae (1 0/65) Acidodontium (5) Anomobryum (6) Brachymenium (3) Bryum (18) Leptobryum (2) Mielichhoferia (6) Orthodontium (2) Pohlia (7) Rhodobryum (5) Schizymenium (11) Callicostaceae (4/4) Callicostellopsis (1) Cyclodicyton (1) Lepidopi/um ( 1) Trachyxiphium (1) Catagoniaceae (111) Catagonium (1) Cryphaeaceae (2/5) Cryphaea (4) Dendrocryphaea ( 1)
273
PARAMOS Daltoniaceae (2114) Daltonia (13) Leskeodon (1) Dicranaceae (17 /67) Aongstroemia (2) Campylopus (37) Chorisodontium (2) Dicrane/la (3) Dicranodontium (!) Dicranoweisia (!) Dicranum (2) Holodontium (1) Ho/omitrium (3) Hygrodicranum (!) Kingiobryum (l) Metzleria (!) Oreoweisia (4) Pi/apagan (3) Rhabdoweisia (1) Sphaerothecium (1) Symb/epharis (3) Ditrichaceae (6/13) Ceratodon (!) Chrysoblaste/la (1) Distichum (1) Ditrichum (5) Pleuridium (3) Tristichium (2) Enca1yptaceae (1/2) Encalypta (2) Entodontaceae (1/1) Enlodan(!) Eustichiaceae (111) Diplostichum (1) Fabroniaceae (1/2) Fabronia (2) Fissidentaceae (1/5) Fissidens (5) Fontina1iaceae (111) Fontinalis (1) Funariaceae (2/12) Entosthodon (8) Funaria (4) Grimmiaceae (4/ 17) Grimmia (7) Ptychomitrium (1) Racomitrium (6) Schistidium (3) Hedwigiaceae (3/7) Braunia (5) Hedwigia (!) Hedwigidium (1) Hylocomiaceae (3/3) Leskeobryum (1) Pleurozium (1) Rhytidium (1) Hypnaceae (7/9) Cariobaeohypnum (1) Ctenidium (1) Herzogie/la ( 1) Hypnum (2) Mittenothamnium (2) Pseudotaxiphy/lum (1) Pylaisie/la (1)
Hypopterygiaceae (111) Hypopterygium (1) Lepyrodontaceae (1/1) Lepyrodon (1) Leskeaceae (2/3) Leskea (2) Leskeade/phus ( 1) Leucodontaceae (111) Leucodon (1) Leucomiaceae (111) Rhychostegiopsis (1) Macromitriaceae (2/ 14) Macrocoma (!) Macromitrium (13) Meesiaceae ( 112) Meesia (2) Meteoriaceae (611 O) Lindigia ( 1) Meteorium (!) Papillaria (3) Pilotriche/la (2) Squamidium (2) Zelometeorium (!) Mniaceae ( 1/l) Plagiomnium (!) Neckeraceae (1/5) Neckera (5) Orthotrichaceae (3/36) Amphidium (1) Orthotrichum (14) Zygodon (21) Phyllogoniaceae (1/2) Phy/logonĂum (2) P1agiotheciaceae (114) Plagiothecium (4) Polytrichaceae ( 6/16) Notoligotrichum (1) 0/igotrichum (1) Pogonatum (4) Polytrichadelphus (6) Polytrichastrum (2) Polytrichum (2) Pottiaceae (19/63) Alonia (1) Aloine/la (3) Anoectangium (1) Barbula (4) Bellibarbula ( 1) Bryoerythrophy/lum (4) Didymodon (8) Erythrophyllastrum (1) Hennedie/la (3) Hymenostylium (1) Leptodontium (16) Mironia (1) Molendoa (1) Pseudocrossidium (2) Pseudosymbe/pharis ( 1) Sagenotortula ( 1) Streptopogon (2) SyntrĂchia (9) Trichostomum (3) Prionodontaceae (1/2) Prionodon (2)
274
MEMOIRS OF THE NEW YORK BOTANICAL GARDEN Racopilaceae ( 112) Racapilum (2) Rhacocarpaceae (1/1) Rhacacarpus ( 1) Rhizogoniaceae (3/3) Leptatheca (1) Pyrrhabryum (!) Rhizaganium (!) Rigodiaceae ( 111) Rigadium (!) Seligeriaceae (1/2) Blindia (2) Sematophyllaceae (4/9) Acraparium (!) Aptychella (1) Aptychapsis (!) Semataphyllum (6) Sphagnaceae ( 1127) Sphagnum (27) Splachnaceae (4/8) Brachymitrian (2) Splachnum (!) Taylaria (4) Tetrapladan (!) Thamnobryaceae (2/5) Paratrichadendran (3) Paratrichum (2) Thuidiaceae (2/5) Cyrta-hypnum (!) Thuidium (4)
Hepatics Anthocerotae Anthocerotaceae ( 111) Phaeaceras (1) Dendrocerotaceae ( 111) Megaceras (!) Hepaticae Acrobolbaceae (4/4)
Acrabalbus ( 1) Lethacalea (1) Marsupidium (!) Tylimanthus (!) Adelanthaceae (1/5) Adelanthus (5) Aneuraceae (2/21) Cryptathallus (1) Riccardia (20) Arnelliaceae (1/3) Gangylanthus (3) Aytoniaceae (111) Asterella (!) Balantiopsidaceae (2/7) Isatachis (6) Ruizanthus (!) Calypogeiaceae (1/3) Calypageia (3) Cephaloziaceae (2/5) Cephalazia (3) Odantaschisma (2) Cepha1oziellaceae (1/6) Cephalaziella (6) Fossombroniaceae (2/4) Austrafassambrania (!) Fassambrania (3)
Geocalycaceae (7/18) Campanacalea (!) Clasmatacalea (!) Heterascyphus (2) Leptascyphus (7) Laphacalea (5) Platycaulis (1)
Pseudacephalaziella (1) Gymnomitriaceae (5/14) Gymnamitrian (4) Marsupe/la (6) Paramamitrian (!) Stephanie/la (2) Stephaniellidium ( 1) Haplomitriaceae (111) Haplómitrium (!) Herbertaceae (3/7,) Herbeftus (5) · Olgantha (!} Triandraphyllum (!) Jubulaceae (1/13) Fru/lania (!3) Jungermanniaceae (11131) Anastraphyllum (8) Andrewsianthus (!) Cryptachila (1) Gymnacaleapsis (!) Jamesaniella (3) Jungermannia (5) Laphanardia (!) Laphazia (4) Nardia (!) Rhadaplagiachila (!) Syzygie/la (5) Lejeuneaceae (16/38) Amphilejeunea (2) Anaplalejeunea (!) Aurealejeunea (3) Blepharalejeunea (2) Brachialejeunea ( 1) Cheilalejeunea (3) Calura (4) Diplasialejeunea (4) Drepanalejeunea (5) Frullanaides (1) Harpalejeunea (5) Leucalejeunea ( 1) Lindigianthus (!) Macralejeunea (!) Micralejeunea (2) Omphalanthus (2) Lepicoleaceae (113) Lepicalea (3) Lepidoziaceae (6/29) Bazzania (!3) Kurzia (2) Lepidazia (8)
Paracramastigum (2) Pseudacephalazia (!) Telaranea (3) Marchantiaceae (2/4) Dumartiera (1) Marchantía (3) Metzgeriaceae ( 1/20) Metzgeria (20) Monocleaceae (1/1) Manaclea (!)
[VOL. 84
Pallaviciniaceae (2/7) Jensenia (2) Symphyagyna (5) Pelliaceae (111) Nateraclada (!) P1agiochi1aceae (2/19) Plagiachila (18) Steereachila ( 1) Pleuroziaceae (111) Pleurazia (!) Porellaceae (112) Parella (2) Pseudolepico1eaceae (2/2) Blepharastama (1) Temnama (!) Radu1aceae (1/6) Radula (6) Ricciaceae (1/2) Riccia (2) Scapaniaceae (2/4) Diplaphyllum (2) Scapania (2) Trichocoleaceae (115) Trichacalea (5)
Vascular Plants Ferns & Fern Allies Adiantaceae (112) Adiantum (2) Asp1eniaceae (2/11) Asplenium (!O) Cystapteris ( 1) Azollaceae (112) Azalla (2) Blechnaceae (1/9) Blechnum (9) Cyatheaceae ( 116) Cyathea (6) Dennstaedtiaceae (3/5) Histiapteris (1) Hypalepis (3)
Pteridium ( 1) Dicksoniaceae (111)
Dicksania ( 1) Dryopteridaceae (5/77) Athyrium (!) Dryapteris (1) Elaphaglassum (65) Palystichum (9) Waadsia (!) Equisetaceae ( 111) Equisetum (!) Gleicheniaceae (113) Gleichenia (3) Hymenophyllaceae ( 1120) Hymenaphyllum (20) Isoetaceae (1/18) Isai!tes (18) Lophosoriaceae ( 111)
Laphasaria ( 1) Lycopodiaceae (3/69) Huperzia (60) Lycapadiella (4) Lycapadium (5)
1999)
275
PARAMOS
Marsi1eaceae ( 1/2) Pilularia (2) Ophioglossaceae (2/4) Botrychium (2) Ophioglossum (2) Plagiogyraceae (111) Plagiogyria (1) Polypodiaceae (14/60) Campy/oneurum (7) Ceradenia (6) Cochlidium (1) Enterosora (1) Grammitis (2) Le/lingeria (4) Melpomene (JI) Micropo/ypodium (!) Niphidium (!) Pec/uma (3)
Pleopeltis (!) Polypodium (12) Terpsichore (9) Zygophlebia (1) Pteridaceae (7/43) Cheilanthes (5) Eriosorus (14) Jamesonia (18) Nephopteris (1)
Pellaea (!) Pityrogramma (2) Pteris (2) Selaginellaceae (1/2) Selaginella (2) Thelypteridaceae (1/13) Thelypteris (13) Vittariaceae (2/2) Radiovittaria ( 1)
Vittaria (!) Gymnosperrns Ephedraceae ( 1/2) Ephedra (2) Angiosperrns Acanthaceae ( 111) Stenandrium ( 1) Alstroemeriaceae ( 1/22) Bomarea (22) Amaranthaceae ( 1/2) Alternanthera (2) Amaryllidaceae (1/2) Stenomesson (2) Apiaceae (15/61) Arracacia (5) Azorella (9) Bowlesia (2) Cotopaxia (2) Daucus (1) Eryngium (2) Hydrocoty/e (12) Lilaeopsis (2) Microp/eura (1) Myrrhidendron (5) Neone/sonia (1) Niphogeton (13) Oreomyrrhis (1)
Ottoa (!) Perissocoeleum (4)
Aquifoliaceae ( 111 O)
Ilex (!O) Araliaceae (2/19)
Oreopanaz (!5) Schejjlera (4) Asclepiadaceae (2/8) Cynanchum (7) Sarcostemma (1) Asteraceae (1011858) Achillea (1)
Achyrocline (1 O) Aequatorium (4) Ageratina (38) Al/oispermum (3) Antennaria (1) Aphanactis (7) Aristeguietia (5) Ascidiogyne (2) Baccharis (45) Badilloa (2) Barnadesia (3) Belloa (7) Bidens (6) Blakiella ( 1) Cabreriella (2) Chaptalia (6) Chersodoma (!) Chevrue/ia ( 1) Chionolaena (2) Chromo/aena (4) Chrysactinium (5) Chuquiraga (3) Cirsium (2) Coespeletia ( 6) Conyza (13) Coreopsis (7) Cotula (2) Critoniopsis (1) Cronquistianthus (4) Cuatrecasasiella ( 1) Diplostephium (70) Dorobaea (2) Erigeron ( 1O) Espeletia (61) Espeletiopsis (21) Face/is (1) Floscaldasia (1) Flosmutisia (!) Freya (1) Galinsoga (2) Gamochaeta (11) Gnaphaliothamnus ( 1) Gnaphalium (19) Grosvenoria (2) Gynoxys (46) Heliopsis (2) Helogyne (!) Hieracium (13) Hinterhubera (8) Hypochaeris (Ji) Iltisia (2) Jalcophila (2) Jaramilloa ( 1) Jungia (6) Laestadia (4) Lasiocepha/us (14)
Libanothamnus (11) Llerasia (3) Loricaria (15) Lourteigia (7) Lucilia (2) Matricaria(!) Mikania (7) Mniodes (!) Monactis (!) Munnozia (7) M ulisia (!O) Myriactis (5) Noticastrum (!) Novenia (1) Oritrophium (15) Oxylobus (1) Pappobolus (8) Paramijlos ( 1) Paranephelius (5) Pentacalia (89) Perezia (3) Philoglossa ( 1) Plagiocheilus (3) Ruouliopsis (2) Ruilopezia (24) Sabazia (4) Scrobicaria (2) Selloa (2) Senecio (69) Sigesbeckia (3) Smallanthus ( 1) Sonchus (2) Stevia (11) Stuckertiella (1) Tagetes (3) Ta/amancalia (1) Taraxacum (3) Trichocline ( 1) Tridax (2) Vasquezia (2) Viguiera (2) Werneria (5) Westoniella (6) Xenophyllum (6) Balanophoraceae (111)
Corynaea (!) Basellaceae (1/1) Tournonia (1) Begoniaceae (1/6) Begonia (6) Berberidaceae (2/32) Berberis (31) Mahonia (1) Bignoniaceae ( 111) Eccremocarpus (1) Boraginaceae (8/13) Cynoglossum (2) Hackelia (3)
Lappula (!) Lithospermum (!) Moritzia ( 1) Myosotis (!) Plagiobothrys (2) Tournefortia (2) Brassicaceae ( 13/71) Brassica ( 1)
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Brayopsis (1) Capsella (1) Cardarnine (11) Crernolobus ( 1) Descurainia (1) Draba{45) Erysirnurn ( 1) Euderna (3) Halirnolobos (1) Lepidiurn (3) ,Rornanschulzia ( 1) Rorippa (1) Brome1iaceae (6/78) Greigia (8) Guzrnania (7) Pitcairnia (4) Puya (48) Racinaea (1) Tillandsia (10) Brunelliaceae (1/8) Brunellia (8) Budd1ejaceae (1/5) Buddleja (5) Cactaceae (111) Opuntia (1) Callitrichaceae (1/4) Callitriche (4) Campanu1aceae (5/48) Centropogon (6) Hypsela (1) Lobelia (3) Lysipornia (27) Siphocarnpylus ( 11) Caprifo1iaceae (1/2) Viburnurn (2) Caryophyllaceae (10/54) Arenaría (15) Cerastiurn (19) Colobanthus ( 1) Dryrnaria ( 6) Paronychia (2) Sagina (1) Scleranthus ( 1) Si/ene (2) Spergularia (1) Stellaria ( 6) Ce1astraceae ( 114) Maytenus (4) Ch1oranthaceae ( 1/6) Hedyosrnurn (6) C1ethraceae ( 115) Clethra (5) C1usiaceae (2/56) Clusia (2) Hypericurn (54) Columelliaceae (1/2) Colurnellia (2) Convolvulaceae (2/2) Cuscuta (1) Dichondra ( 1) Coriariaceae ( 111) Cariaría (1) Crassu1aceae (3/9) Crassula (4) Echeveria (4) Villadia (1)
Cunoniaceae (1114) Weinrnannia (14) Cyperaceae (8/70) Bulbostylis ( 1) Carex (31) Cyperus (3) Eleocharis ( 1O) Jsolepis (3) Oreobolus ( 5) Rhynchospora (10) Uncinia (7) Desfontainiaceae (112) Desfontainia (2) Dioscoreaceae ( 111) Dioscorea (1) Droseraceae (1/1) Drosera (1) · E1aeocarpaceae.(l/1) Val/ea (1) E1atinaceae (1/6) Elatine (6) Eremo1epidaceae (111) Antidaphne (1) Ericaceae (16/79) Bejaria (5) Cavendishia (3) Ceratosterna (2) Cornarostaphylis ( 1) Demosthenesia (2) Disterigrna (8) Gaultheria (19) Gaylussacia (1) Macleania (2) Pernettya (2) Plutarchia (1 O) Psarnrnisia (4) Sernirarnisia (2) Themistoclesia (8) Thibaudia (4) Vacciniurn (6) Eriocaulaceae (3/28) Eriocaulon (1) Paepalanthus (26) Syngonanthus (1) Euphorbiaceae (3/3) Charnaesyce ( 1) Dysopsis (1) Euphorbia (1) Fabaceae (9/76) Astragalus (3) Cologania (1) Dalea (3) Lathyrus (2) Lupinus (56) Medicago (2) Otholobiurn (3) Trifoliurn (4) Vicia (2) Garryaceae (111) Garrya (1) Gentianaceae (4/93) Gentiana (3) Gentianella (48) Halenia (39) Macrocarpaea (3)
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Geraniaceae (2/45) Erodium (2) Geranium (43) Gesneriaceae (3/4) Alloplectus (2) Colurnnea ( 1) Heppiella (1) Grossu1ariaceae (2/13) Escallonia (1) Ribes (12) Ha1oragaceae (2/13) Gunnera (11) Myriophyllurn (2) Hydrocharitaceae (112) Elodea (2) Hydrophyllaceae (111) Phacelia (1) Hypoxidaceae (1/2) Hypoxis (2) Iridaceae (2115) Orthrosanthus (4) Sisyrinchium (11) Juncaceae (4/24) Distichia (2) Juncus (16) Luzula (5) Rostkovia (1) Juncaginaceae (1/1) Lilaea (1) Lamiaceae ( 6/3 7) Lepechinia (4) Minthostachys (1) Prunella (1) Salvia (15) Sature) a (9) Stachys (7) Lauraceae (1/2) Persea (2) Lemnaceae ( 111) Lernna (1) Lentibu1ariaceae (2/6) Pinguicula (5) Utricularia (1) Liliaceae (3/5) Echeandia (1) Excremis (1) Jsidrogalvia (3) Loasaceae (2/2) Caiophora (1) Loasa (1) Loranthaceae (3/6) Aetanthus (3) Gaiadendron (1) Tristerix (2) Lythraceae (1/2) Cuphaea (2) Malvaceae (2/17) Acaulirnalva (1 O) Nototriche (7) Me1astomataceae (9/1 07) Axinaea (2) Brachyotum (22) Bucquetia (2) Castratella (2) Chaetolepis (8) · Meriania (1)
1999]
Miconia (54) Monochaetum (12) Tibouchina (4) Myricaceae (1/3) Myrica (3) Myrsinaceae (3/J O) Cybianthus (4) Geissanthus (2) Myrsina (4) Myrtaceae (4/7) Eugenia (J) Myrcianthes (2) Myrteola (3) Ugni (!) Nyctaginaceae (1/2) Colignonia (2) Onagraceae (3/27) Epilobium (J) Fuchsia (22) Oenothera (4) Orchidaceae (25/J52) Aa (9) Altensteinia (3) Barbase/la (1) Brachionidium (2) Cranichis (2) Elleanthus (8) Epidendrum (27) Gomphichis (13) Habenaria (J) Lepan/hes ( 11) Malaxis (1) Masdevallia (5) Maxillaria (4) Myrosmodes (6) Odontoglossum (9) Oncidium (2) Pachyphyllum (7) Pleurothallis ( J5) Pterichis (8) Salpistele (1) Stelis (8) Stenorrhynchos (1) Telipogon (4) Ticoglossum (1) Trichosalpinx (3) Oxa1idaceae (1/1 O) Oxalis (10) Passifloraceae (1/16) Passiflora (16) Phyto1accaceae (114) Phytolacca (4) Piperaceae (2/19) Peperomia (16) Piper (3) P1antaginaceae (1/1 O) Plantago (1 O) Poaceae (41122 7) Aciachne (3) Aegopogon (1) Agrostis (24) Alopecurus (1) Axthoxanthum (1) Aphanelytrum (1) Aristida ( J)
PARAMOS
Arthrostylidium (!) Aulonemia (4) Bothriochloa ( 1) Brachypodium (1) Eriza (J) Bromus (6) Calamagrostis (36) Chusquea ( 1O) Cinna (J) Cortaderia (5) Dactylis (J) Danthonia (1) Dissanthelium (1) Elymus (J) Festuca (38) Hierochloe (2) Holcus (1) Melica (J) Muhlenbergia (1 1) Nassella ( 6) Neurolepis (JO) Ortachne ( 1) Paspalum (7) Phalaris (2) Piptochaetium (4) Poa (20) Poidium (1) Polypogon (2) Schizachyrium (1) Sporobolus (2) Stipa (4) Triniochloa (2) Trisetum ( 6) Vulpia (4) Po1yga1aceae (1/1 7) Monnina (J7) Po1ygonaceae (3/11) Muehlenbeckia (4) Polygonum (1) Rumex (6) Portu1acaceae (2/6) Calandrinia (3) M antia (3) Potamogetonaceae (2/4) Potamogeton (3) Stuckenia (!) Primu1aceae (111) Lysimachia (1) Proteaceae (2/2) Oreocallis (1) Roupala (J) Ranuncu1aceae (6/2J) Anemone (1) Caltha (1) Laccopetalum (1) Oreithales ( J) Ranunculus (1 6) Thalictrum (!) Rhamnaceae (l/2) Rhamnus (2) Rosaceae ( J 0/77) Acaena (6) Aphanes (1) Fragaria (1) Geum (1)
277
Hesperomeles (7) Lachemilla (34) Polylepis (JO) Poten tilla (2) Rubus (J4) Sericotheca (J) Rubiaceae (6/30) Arcytophyllum (J4) BarrerĂa ( 1) Galium (10) Manettia (2) Nertera (2) Sherardia ( 1) Saxifragaceae (111) SaxĂfraga ( J) ScrophuJariaceae (J4/J44) Alonsoa (8) Aragoa (17) Bartsia (22) Calceolaria (65) Castilleja (1 5) Digitalis ( 1) Gratiola (!) Lamourouxia (2) Limos ella (4) Mimulus (1) Ourisia (2) Pedicularis (1) Sibthorpia (1) Veronica (4) So1anaceae (8/58) Cestrum (4) Deprea (J) Jaltomata (2) Nicotiana (1) Nierembergia (2) Salpichroa (3) Saracha (2) Solanum (43) Symp1ocaceae (111 O) Symplocos (JO) Theaceae ( 112) Ternstroemia (2) Tropaeo1aceae (1/J) Tropaeolum (1) Urticaceae (3/8) Parietaria ( J) Pilea (3) Urtica (4) Va1erianaceae (2/55) Belonanthus ( 1) Valeriana (54) Verbenaceae (3/5) Hierobotana (J) Lippia (1) Verbena (3) Vio1aceae (2/1 6) Hybanthus (1) Viola (15) Viscaceae (1110) Dendrophthora ( 1O) W interaceae ( 111) Drimys (1) Xyridaceae ( 111) Xyris (1)
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MEMOIRS OF THE NEW YORK BOTAN! CAL GARDEN
Appendix 2: Nomenclatural Novelties
Hepaticae Cheilolejeunea choachina (Gottsche) Gradst., see p. 69 Isotachis lopezii (R.M.Schust.) Gradst., see p. 66
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