00 oliveira fm 7/31/02 8:11 AM Page i The Cerrados of Brazil 00 oliveira fm 7/31/02 8:11 AM Page ii 00 oliveira fm 7/31/02 8:11 AM Page iii The Cerrados of Brazil Ecology and Natural History of a Neotropical Savanna Editors Paulo S. Oliveira Robert J. Marquis Columbia University Press New York 00 oliveira fm 7/31/02 8:11 AM Page iv Columbia University Press Publishers Since 1893 New York Chichester, West Sussex © 2002 Columbia University Press All rights reserved Library of Congress Cataloging-in-Publication Data The cerrados of Brazil : ecology and natural history of a neotropical savanna / Paulo S. Oliveira and Robert J. Marquis. p. cm. Includes bibliographical references. ISBN 0-231-12042-7 (cloth : alk. paper)—ISBN 0-231-12043-5 (pbk. : alk. paper) 1. Cerrado ecology—Brazil. I. Oliveira, Paulo S., 1957– II. Marquis, Robert J., 1953– QH117 .C52 2002 577.4'8'0981—dc21 2002022739 Columbia University Press books are printed on permanent and durable acid-free paper. Printed in the United States of America c 10 9 8 7 6 5 4 3 2 1 p 10 9 8 7 6 5 4 3 2 1 00 oliveira fm 7/31/02 8:11 AM Page v Contents Preface vii 1 Introduction: Development of Research in the Cerrados 1 Paulo S. Oliveira and Robert J. Marquis I Historical Framework and the Abiotic Environment 2 Relation of Soils and Geomorphic Surfaces in the Brazilian Cerrado 13 Paulo E. F. Motta, Nilton Curi, and Donald P. Franzmeier 3 Late Quaternary History and Evolution of the Cerrados as Revealed by Palynological Records 33 Marie-Pierre Ledru 4 The Fire Factor 51 Heloisa S. Miranda, Mercedes M. C. Bustamante, and Antonio C. Miranda 5 Past and Current Human Occupation, and Land Use 69 Carlos A. Klink and Adriana G. Moreira II The Plant Community: Composition, Dynamics, and Life History 6 Vegetation Physiognomies and Woody Flora of the Cerrado Biome 91 Ary T. Oliveira-Filho and James A. Ratter 7 Herbaceous Plant Communities 121 Tarciso S. Filgueiras 8 Patterns and Dynamics of Plant Populations 140 Raimundo P. B. Henriques and John D. Hay 9 The Role of Fire in Population Dynamics of Woody Plants 159 William A. Hoffmann and Adriana G. Moreira v 00 oliveira fm 7/31/02 8:11 AM Page vi vi contents 10 Ecophysiology of Woody Plants 178 Augusto C. Franco III The Animal Community: Diversity and Biogeography 11 Lepidoptera in the Cerrado Landscape and the Conservation of Vegetation, Soil, and Topographical Mosaics 201 Keith S. Brown Jr. and David R. Gifford 12 The Character and Dynamics of the Cerrado Herpetofauna 223 Guarino R. Colli, Rogério P. Bastos, and Alexandre F. B. Araujo 13 The Avifauna: Ecology, Biogeography, and Behavior 242 Regina H. F. Macedo 14 The Cerrado Mammals: Diversity, Ecology, and Natural History 266 Jader Marinho-Filho, Flávio H. G. Rodrigues, and Keila M. Juarez IV Insect-Plant Interactions 15 Ant Foraging on Plant Foliage: Contrasting Effects on the Behavioral Ecology of Insect Herbivores 287 Paulo S. Oliveira, André V. L. Freitas, and Kleber Del-Claro 16 Interactions Among Cerrado Plants and Their Herbivores: Unique or Typical? 306 Robert J. Marquis, Helena C. Morais, and Ivone R. Diniz 17 Pollination and Reproductive Biology in Cerrado Plant Communities 329 Paulo E. Oliveira and Peter E. Gibbs V The Conservation of the Cerrados 18 Biodiversity and Conservation Priorities in the Cerrado Region 351 Roberto B. Cavalcanti and Carlos A. Joly Contributors 369 Index 373 00 oliveira fm 7/31/02 8:11 AM Page vii Preface This is a book about the Cerrado Biome, a major Brazil- ian savanna-like ecosystem for which no such summary exists. Biologists outside Brazil know little about the cerrados, despite the fact that the biome covers approximately 22% of the country’s surface area, or 2 mil- lion km2. Even though much of the attention of conservationists has focused on rainforests such as the Amazon and Atlantic forests, the cer- rados are currently one the most threatened biomes of South America due to the rapid expansion of agriculture. Nearly 50% of the cerrado region is currently under direct human use, and about 35% of its total natural cover has been converted into planted pastures and crops. The average annual rate of land clearing in the cerrados during 1970–1975 was nearly twice the estimated deforestation rate of the Amazon forest during 1978–1988. Overall biodiversity for the Cerrado Biome, including all its physiognomic forms, is estimated at 160,000 species of plants, animals, and fungi. Endemicity of cerrado higher plants has recently been esti- mated at 4,400 species, representing 1.5% of the world’s total vascular plant species. Endemic vertebrates range from 3% (birds) to 28% (am- phibians) of the species recorded. The cerrados are also unique in that they serve as corridors for species inhabiting neighboring biomes such as the Amazonian and Atlantic rainforests. For example, although endemicity is low among birds, 90% of the species breed in the cerrado region. Given their geographic extent, it is surprising that the cerrados remain largely ignored at the international level. Because of the threatened status and rich biodiversity of this Neotropical savanna, and the lack of familiarity with cerrado ecosystems at the international level, a volume that compiles the known natural history, ecology, and biogeography of this biome is extremely timely. This is perhaps the first volume in English covering a tropical ecosys- tem in which the vast majority of the contributors are from the region in question. The foreign exceptions include scientists that are very familiar with the cerrados and have long-lasting collaborations with Brazilian researchers. The volume is broad in scope and raises relevant ecological questions from a diversity of fields, indicating areas in which additional vii 00 oliveira fm 7/31/02 8:11 AM Page viii viii preface research is needed. Such a wide thematic approach should provide the international audience with a broad ecological framework for under- standing the cerrado savanna. The editors hope that such a book will make an important contribution for ecology, and for tropical biology in particular, stimulating future research in the cerrados. The idea of preparing a book summarizing research on cerrado biol- ogy arose in 1997 in San José, Costa Rica, during a most exciting meet- ing of the Association for Tropical Biology. As the book project devel- oped, a number of people helped us shape the scope of the volume, establishing the main research areas to be covered, adjusting chapter con- tents, and writing the book proposal. At the early stages we have bene- fited greatly from the encouragement as well as the technical and editor- ial experience of Susan E. Abrams of the University of Chicago Press and Peter W. Price of Northern Arizona University. Helpful suggestions were also given by Keith S. Brown, William A. Hoffmann, Regina Macedo, Ary T. Oliveira-Filho, and Guy Theraulaz. Humberto Dutra helped with the preparation of the book index, and Glauco Machado and André Freitas helped with the scanning and printing of the figures. Mailing costs were covered in part by the Ecology Graduate Program of the Universidade Estadual de Campinas. Each chapter was substantially improved by the comments and sug- gestions of external reviewers. They include Steve Archer, John A. Barone, Kamaljit S. Bawa, John G. Blake, Keith S. Brown, Ray B. Bryant, Phyllis D. Coley, Philip J. DeVries, Peter E. Gibbs, Guillermo Goldstein, Gary S. Hartshorn, W. Ronald Heyer, Peter Kershaw, W. John Kress, Thomas H. Kunz, Diana Lieberman, Arício X. Linhares, Vera Markgraf, Ernesto Medina, Daniel C. Nepstad, Ary T. Oliveira-Filho, James L. Patton, A. Townsend Peterson, Ghillean T. Prance, Peter W. Price, James A. Ratter, José F. Ribeiro, Juan F. Silva, Robert B. Srygley, and Laurie J. Vitt. We appreciate the time they took to give critical reviews. Finally, we thank Science Editor Holly Hodder and Assistant Editor Jonathan Slutsky, formerly of Columbia University Press, for their initial encouragement and advice on the development of this project. Current Assistant Editor Alessandro Angelini helped at the final stage of the edit- ing process, and Diana Senechal copyedited the entire manuscript. We are especially grateful to Julie S. Denslow and Lucinda A. McDade, review- ers of the book proposal for Columbia University Press, for their careful and constructive suggestions concerning the initial book project. Paulo S. Oliveira Robert J. Marquis 00 oliveira fm 7/31/02 8:11 AM Page ix The Cerrados of Brazil 00 oliveira fm 7/31/02 8:11 AM Page x 01 oliveira ch 1-2 7/31/02 8:12 AM Page 1 1 Introduction: Development of Research in the Cerrados Paulo S. Oliveira and Robert J. Marquis The first detailed account of the Brazilian cerrados was provided by Danish botanist Eugene Warming (1892) in the book Lagoa Santa, in which he describes the main features of the cerrado veg- etation in the state of Minas Gerais. Since the publication of Warming’s book a number of descriptive studies from several cerrado regions in Brazil have been published. The vast majority of this literature is in Por- tuguese and oriented mostly toward botanical aspects of the cerrado. The studies can be roughly categorized into two major groups: (1) Surveys of woody floras, frequently providing also the general physiognomic char- acteristics of the vegetation (thorough reviews of this literature are given by Eiten 1972; Goodland and Ferri 1979). (2) Studies on plant ecophysi- ology focusing particularly on mineral nutrition, fire, and water economy at the plant-soil and plant-atmosphere levels; and on how these factors can account for the characteristic xeromorphic aspect of cerrado woody plants (extensive lists of these studies are given by Labouriau 1966; Ferri 1977; Goodland and Ferri 1979).