570.5 ILL 59 " 199 0- A Taxonomic mv of the Palm Subtribe Attaleinae (Tribe Cocoeae) SIDiNLV i\ GLASSMAN UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN BIOLOGY 2891 etc 2 8 Digitized by tine Internet Arcinive in 2011 witii funding from University of Illinois Urbana-Champaign http://www.archive.org/details/taxonomictreatme59glas A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) SIDNEY F. GLASSMAN ILLINOIS BIOLOGICAL MONOGRAPHS 59 UNIVERSITY OF ILLINOIS PRESS Urbana and Chicago Illinois Biological Monographs Committee David S. Seigler, chair Daniel B. Blake Joseph V. Maddox Lawrence M. Page © 1999 by the Board of Trustees of the University of Illinois Manufactured in the United States of America P 5 4 3 2 1 This edition was digitally printed. © This book is printed on acid-free paper. Library of Congress Cataloging-in-Publication Data Classman, Sidney F. A taxonomic treatment of the palm subtribe Attaleinae (tribe Cocoeae) / Sidney F. Classman. p. cm. — (Illinois biological monographs ; 59) Includes bibliographical references (p. ) and index. ISBN 0-252-06786-X (pbk. : acid-free paper) 1. Palms—Classification. 2. Palm.s—Classification —South America. I. Title. II. Series. QK495.P17G548 1999 584'.5—dc21 98-58105 CIP Abstract Detailed taxonomic treatments of all genera in the subtribe AttaleinaesLve included in the present study. Attalea contains 21 species; 2 interspecific hybrids, A. x piassabossu and A. x voeksii; and 1 intergeneric hybrid, x Attabignya minarum. Orbignya has 1 1 species; 1 interspecific hybrid, O. x teixeirana; 2 intergeneric hybrids, x Attabignya minarum and x Maximbignya dahlgreniana; 1 putative intergeneric hybrid, Ynesa colenda; and 1 unde- scribed putative intergeneric hybrid. Scheelea contains 31 species and 2 undescribed putative intergeneric hybrids. Maximiliana conX2iins 1 species with 1 intergeneric hybrid, x Maximbignya dahlgreniana, and 1 undescribed putative intergeneric hybrid. A total of 13 new species are described here {Attalea, 3, Orbignya, 1 , and Scheelea, 9) , as well as 3 new combinations and name transfers. The main emphasis of this treatment is taxonomic, but other system- atic aspects have been considered, such as leaf anatomy and chemotax- onomy as well as geographical and ecological distribution. Evolutionary relationships among all four genera, as well as alliances among species within each genus (including a tentative division into infrageneric catego- ries) , are followed by a section on geographical distribution of species within each genus, a description of each genus, and keys to their species. Taxonomic treatments of all four genera and their individual species in- clude original as well as other pertinent publications, complete synonymy, designation of types, a detailed morphological description, a list of speci- mens examined, distribution and habitat, vernacular names, economic importance, a discussion of evolutionary relationships, and nomenclatural problems. A total of 10 tables, 201 illustrations, and 17 distribution maps are also included. Illustration and map numbers are listed as Figs, in bold type for each species. Lists of doubtful and excluded species for each genus are also incorpo- rated into the text, as are acknowledgements, a list of references, and an alphabetical index to names of all taxa in the subtribe Attaleinae. Cladistic studies are not included in the present treatment due to in- complete diagnostic morphological data for a number of taxa. At this juncture I should point out that this work does not represent a conven- tional monograph or revision. However, the total amount of information contained in it, and found nowhere else, forms a basis for future study that should lead to a more complete monographic treatment. Therefore, the main purpose of this study is to lay the groundwork to stimulate further research (including the collection of additional specimens) of this com- plex group of palms. Acknowledgments I am indebted to the National Science Foundation for three grants (GB6899, GB3737, and BMS7509779) and to the Research Board and the Department of Biological Sciences at the University of Illinois at Chicago for additional assistance. These grants enabled me to take field trips to Brazil; visit herbaria in Europe, South America, and the United States; and study cultivated palms of various genera in the subtribe Attaleinae at Fairchild Tropical Garden in Miami, Florida. I would like to thank J. B. Harborne and C. Williams, both at the University of Reading, England, for carrying out chemotaxonomic studies on flavonoids; the late Timothy Plowman of the Field Museum for numerous valuable suggestions; and the late Rolf Singer of the Field Museum for revising the Latin descrip- tions of new species. Special thanks go to Larry Noblick of Montgomery Botanical Center for important field information on the distribution of Attalea in Bahia; to Margaret Kleist andJoan Cureton of UIC for prepara- tion of the original manuscript; to my son Bob for his computer exper- tise and excellent editorial assistance; to my wife, Ida, for continuous sup- port in the preparation of this manuscript; to Marlene Werner of the Field Museum and Janice Rajecki of UIC for preparation of the illustrations; to several people at the Instituto Botanica, Sao Paulo, especially A. Milanez and D. Vital, for making available to me transportation, guides, drivers, and herbarium facilities during my trip to Brazil in 1976; to the late Pedrito Silva of CEPLAC in Salvador for facilitating the shipment of palm speci- mens to the United States and for useful information on the disposition of Gregorio Bondar's palm collections; to Leticia Faria of the Federal University of Salvador, Edna Olivera of EMBRAPA in Salvador, Luis Silva of CEPEC in Itabuna, and Luciano Lima of CEPLAC in Petrolina for making available transportation, drivers, and herbarium facilities (all of which was coordinated by Larry Noblick) during our trip to Bahia, Bra- zil, in 1988. 1 am also grateful to the curators of the herbaria cited for the privilege of studying their specimens. Herbaria Cited A Arnold Arboretum, Cambridge, Mass. ALCB Herbario Alexandre Leal Costa, Institute de Biologia, Salva- dor, Brazil B Botanisches Museum, Berlin-Dahlem BAH Herbario Antonio Nonata Marques, Salvador, Brazil BH Liberty Hyde Bailey Hortorium, Ithaca, N.Y. BM British Museum, London BR Herbarium, Jardin Botanique National de Belgique, Meise, Belgium C Botanical Museum and Herbarium, Copenhagen CAS Herbarium, California Academy of Sciences, San Francisco CAY Herbarie Centre ORSTOM, Cayenne, French Guiana CEN Herbario CENARGEN/EMBRAPA, Brasilia, D.F, Brazil CEPEC Herbario de Centro de Pesquisas do Cacau, Itabuna, Brazil COL Institute de Ciencias Naturales, Bogota CPATSA Herbario, Centro de Pesquisas Tropica, Petrolina, Brazil F Field Museum of Natural History, Chicago FI Herbarium Universitatis Florentinae, Florence, Italy FTG Fairchild Tropical Garden, Miami G Conservatoire et Jardin Botaniques, Geneva GH Gray Herbarium of Harvard University, Cambridge, Mass. HUEFS Herbario Universidade Estual, Feira Santana, Brazil IPA Institute de Pesquisas Agronomicas, Dois Irmaos, Recife, Bra- zil K Royal Botanic Gardens, Kew, Great Britain LE Botanical Institute, Academy of Sciences of the USSR, Lenin- grad M Botanische Staatssamlung, Munich MG Museu Paraense Emilio Goeldi, Belem, Brazil MICH Herbarium, University of Michigan, Ann Arbor MO Missouri Botanical Garden, St. Louis NY New York Botanical Garden, Bronx, N.Y P Museum National d'Histoire Naturelle, Paris PMA Herbario, Universidad de Panama, Panama R Museu Nacional, Rio de Janeiro RB Jardim Botanico, Rio de Janeiro S Naturhistoriska Riksmuseum, Stockholm SP Instituto de Botanica, Sao Paulo U Botanical Museum and Herbarium, Utrecht UB Universidade de Brasilia, D.F. US United States National Museum, Washington, D.C. USM Herbario San Marcos, Lima VEN Instituto Botanico, Caracas 71 Contents Abstract v Acknowledgments vii Herbaria Cited ix Introduction 1 History of Subtribe Attaleinae 3 Systematic Studies of Subtribe Attaleinae 6 Genus Attalea 1 1 Geographic Distribution of Genus Attalea 1 Relationships within Genus Attalea 13 Outline of Tentative Division of Attalea into Infrageneric Categories 16 Taxonomic Treatment of Genus Attalea 1 Key to Species of Attalea 20 Taxonomic Treatment of Species of Attalea 24 Doubtful and Uncertain Species of Attalea 67 Genus Orbignya 73 Geographic Distribution of Genus Orbignya 73 Relationships within Genus Orbignya 75 Outline of Tentative Division of Orbignyainto Infrageneric Categories 77 Taxonomic Treatment of Genus Orbignya 78 Key to Species and Hybrids of Orbignya 80 Taxonomic Treatment of Species of Orbignya 82 Doubtful and Uncertain Species of Orbignya 107 Genus Scheelea 115 Geographic Distribution of Genus Sc^e^fefl 115 Relationships within Genus ScAe^fea 117 Outline of Tentative Division of Scheelea into Infrageneric Categories 119 Taxonomic Treatment of Genus Scheelea 120 Key to Species of Scheelea 123 Taxonomic Treatment of Species of Scheelea 128 Doubtful and Uncertain Species of Scheelea 177 Genus Maximiliana Geographic Distribution of Genus Maximiliana 181 Taxonomic Treatment of Species of Maximiliana 182 Doubtful and Uncertain Species oi Maximiliana 185 Hybrids in the Attaleinae 187 Hybrids of Attalea 1 88 Hybrids of Orbignya 197 Hybrids of Scheelea 202 Hybrids of Maximiliana 203 Excluded Species of Attalea 205 Excluded Species of Maximiliana 207 Glossary 209 References 213 Attalea Figures 223 Attalea Pinnae Cross Sections