losos_fm.qxd 4/11/09 9:52 AM Page i LIZARDS IN AN EVOLUTIONARY TREE losos_fm.qxd 4/11/09 9:52 AM Page ii ORGANISMS AND ENVIRONMENTS Harry W. Greene, Consulting Editor 1. The View from Bald Hill: Thirty Years in an Arizona Grassland, by Carl E. Bock and Jane H. Bock 2. Tupai: A Field Study of Bornean Treeshrews,by Louise H. Emmons 3. Singing the Turtles to Sea: The Comcáac (Seri) Art and Science of Reptiles, by Gary Paul Nabhan 4. Amphibians and Reptiles of Baja California, Including Its Pacific Islands and the Islands in the Sea of Cortés, by L. Lee Grismer 5. Lizards: Windows to the Evolution of Diversity, by Eric R. Pianka and Laurie J. Vitt 6. American Bison: A Natural History,by Dale F. Lott 7. A Bat Man in the Tropics: Chasing El Duende,by Theodore H. Fleming 8. Twilight of the Mammoths: Ice Age Extinctions and the Rewilding of America, by Paul S. Martin 9. Biology of Gila Monsters and Beaded Lizards,byDaniel D. Beck 10. Lizards in an Evolutionary Tree: Ecology and Adaptive Radiation of Anoles, by Jonathan B. Losos losos_fm.qxd 4/11/09 9:52 AM Page iii LIZARDS IN AN EVOLUTIONARY TREE Ecology and Adaptive Radiation of Anoles Jonathan B. Losos UNIVERSITY OF CALIFORNIA PRESS Berkeley Los Angeles London losos_fm.qxd 4/13/09 8:18 PM Page iv the publisher and author gratefully acknowledge the generous contribution to this book provided by museum of comparative zoology, harvard university University of California Press, one of the most distinguished university presses in the United States, enriches lives around the world by advancing scholarship in the humanities, social sciences, and natural sciences. Its activi- ties are supported by the UC Press Foundation and by philanthropic contri- butions from individuals and institutions. For more information, visit www.ucpress.edu. Organisms and Environments, No. 10 University of California Press Berkeley and Los Angeles, California University of California Press, Ltd. London, England © 2009 by the Regents of the University of California All photographs are property of the author unless otherwise indicated. Library of Congress Cataloging-in-Publication Data Losos, Jonathan B. Lizards in an evolutionary tree :ecology and adaptive radiation of anoles / Jonathan B. Losos. p. cm. Includes bibliographical references and index. ISBN 978-0-520-25591-3 (cloth : alk. paper) 1. Anoles—Evolution. 2. Anoles—Ecology. 3. Anoles—Adaptation. I. Title. QL666.L268L67 2009 597.95'48138—dc22 2008027796 Manufactured in Singapore 16 15 14 13 12 11 10 09 10 9 8 7 6 5 4 3 2 1 The paper used in this publication meets the minimum requirements of ANSI/NISO Z39.48-1992 (R 1997) (Permanence of Paper). Cover illustration: Anolis monticola.Photo copyright Eladeo Fernández. losos_fm.qxd 4/11/09 9:52 AM Page v To my wife, Melissa Losos, my parents, Carolyn and Joseph Losos, and my sisters, Carol, Elizabeth, and Louise Losos, for their love and support throughout my life losos_fm.qxd 4/11/09 9:52 AM Page vi The Page Left Intentionally Blank losos_fm.qxd 4/11/09 9:52 AM Page vii CONTENTS Foreword by Harry W. Greene • ix Acknowledgments • xiii Prologue: The Case for Anolis • xvii 1. Evolutionary Biology as a Historical Science • 1 2. Meet the Anoles! • 11 3. Five Anole Faunas, Part One: Greater Antillean Ecomorphs • 29 4. Five Anole Faunas, Part Two: The Other Four • 59 5. Phylogenetics, Evolutionary Inference, and Anole Relationships • 81 6. Phylogenetic Perspective on the Timing and Biogeography of Anole Evolution • 99 7. Evolution of Ecomorphological Diversity • 113 8. Cradle to Grave: Anole Life History and Population Biology • 135 9. Social Behavior, Sexual Selection, and Sexual Dimorphism • 161 10. Habitat Use • 189 11. Ecology and Adaptive Radiation • 205 vii losos_fm.qxd 4/11/09 9:52 AM Page viii 12. Natural Selection and Microevolution • 233 13. Form, Function, and Adaptive Radiation • 257 14. Speciation and Geographic Differentiation • 291 15. The Evolution of an Adaptive Radiation • 317 16. The Five Faunas Reconsidered • 351 17. Are Anoles Special, and If So, Why? • 383 Afterword • 411 References • 421 Index • 495 viii • CONTENTS losos_fm.qxd 4/11/09 9:52 AM Page ix FOREWORD Lizards in an Evolutionary Tree: Ecology and Adaptive Radiation of Anoles is the tenth volume in the University of California Press’s series on organisms and environments, whose unifying themes are the diversity of plants and animals, the ways they interact with each other and with their surroundings, and the implications of those relationships for science and society. We seek books that promote unusual, even unexpected connec- tions among seemingly disparate topics, distinguished by the talents and perspectives of their authors. Previous volumes have spanned topics as diverse as grassland ecology and bison behavior, but none has encompassed the breadth and depth of scholarly coverage achieved here. Jonathan Losos chronicles the details and historical underpinnings of an extraordi- nary natural legacy, the adaptive radiation of almost four hundred species of very special lizards. Thanks to their unusual diversity, abundance, and tractability, anoles have played central roles in several scientific disciplines, including physiological and commu- nity ecology, functional morphology, biogeography and molecular evolution. Losos has synthesized anole biology in lively prose, based on his own extensive studies and thou- sands of publications by an army of researchers over the past century. From a conceptual perspective, this book explores the cutting edges of evolutionary biology and ecology, our search for patterns and causal explanations for biodiversity. Why are there more species in some places than others, and what drives diversification? How do individuals interact with others of their species? Is competition among species important? And what will be the fate of anoles on our rapidly changing planet? ix
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