The Ecology & Behavior of Amphibians a frog-catcher The lower sort of people in China are not delicate in their eating; they do not scruple to feed heartily upon animals which die a natural death, as is noticed by several voyagers. The poor eat frogs and rats, dried hams of which latter are exposed to sale in the streets; and the mid- dlingsort think a young dog no bad food .... They have a method of taking frogs in China, during the night, by means of fire, which is carried in a wire net, as reproduced in this plate. —George Henry Mason, The Costume of China(London: W. Miller, 1800). Traditional frog catchers in China probably had detailed knowledge of the natural his- tory of the animals they sought as food. Here the image of the frog-catcher represents the many biologists who have ventured into the field, often at night, to study the ecology and behavior of amphibians. This print is dated May 4, 1799, from a drawing by a Chinese artist, Pu-Qua, of Canton. From the author’s collection. The Ecology & Behavior of Amphibians Kentwood D. Wells The University of Chicago Press Chicago and London kentwood d. wells is professor in the Department of Ecology and Evolutionary Biology at the University of Connecticut The University of Chicago Press, Chicago 60637 The University of Chicago Press, Ltd., London © 2007 by The University of Chicago All rights reserved. Published 2007 Printed in the United States of America 16 15 14 13 12 11 10 09 08 07 1 2 3 4 5 isbn-13: 978-0-226-89334-1 (cloth) isbn-13: 978-0-226-89335-8 (paper) isbn-10: 0-226-89334-0 (cloth) isbn-10: 0-226-89335-9 (paper) Library of Congress Cataloging-in-Publication Data Wells, Kentwood David, 1948– The Ecology and behavior of amphibians / Kentwood D. Wells. p. cm. Includes bibliographical references and index. isbn-13: 978-0-226-89334-1 (cloth : alk. paper) isbn-10: 0-226-89334-0 (cloth : alk. paper) isbn-13: 978-0-226-89335-8 (pbk. : alk. paper) isbn-10: 0-226-89335-9 (pbk. : alk. paper) 1. Amphibians—Ecology. 2. Amphibians—Behavior. I. Title. ql641.w35 2007 597.8—dc22 2007001687 oThe paper used in this publication meets the minimum requirements of the American National Standard for Information Sciences—Permanence of Paper for Printed Library Materials, ansiz39.48-1992. This book is for Marta, Camila, and Gabriela Contents Preface ix 1 Phylogeny, Classification, and Morphological Evolution 1 2 Water Relations 82 3 Temperature Relations 122 4 Respiration 157 5 Metabolism and Energetics 184 6 Movements and Orientation 230 7 Anuran Vocal Communication 268 8 Mating Systems and Sexual Selection in Anurans 338 9 Communication and Social Behavior of Urodeles and Caecilians 403 10 The Natural History of Amphibian Reproduction 451 11 Parental Care 516 12 The Ecology and Behavior of Amphibian Larvae 557 13 Complex Life Cycles and the Ecology of Amphibian Metamorphosis 599 14 Amphibians and Their Predators 645 15 The Ecology of Amphibian Communities 729 16 Conservation of Amphibians 784 References 857 Index 1085 Preface The train from Balboa Station moved slowly at understatement. Indeed, at times the project seemed to take first, making its way past the warehouses, military on a life of its own, like one of those never-ending scholarly bases, and red-tiled roofs of Canal Zone houses, endeavors usually associated with the humanities, such as past the locks of Miraflores and Pedro Miguel, and along- editing the correspondence of Charles Darwin or writing side the Gaillard Cut. There it picked up speed, traveling the definitive multivolume biography of Herman Melville. parallel to the Panama Canal and eventually reaching Much has changed in the intervening years, both in my life stretches of tropical rainforest, dotted with small farms and and in the study of amphibian ecology and behavior. I began banana fields. As I gazed out the window at the ponds and the book as a young assistant professor and now find myself lakes that lay alongside the tracks, I thought about all the one of the most senior members of my department. I got frogs that must be out there, feeding, calling, defending ter- married, raised two young daughters, bought and remod- ritories, laying eggs, avoiding snakes, and otherwise going eled a house, taught classes, supervised graduate students, about their lives. I couldn’t see them, of course, nor could I conducted field research, and served on more departmental hear them at night over the din and clatter of the train, but and university committees than I can remember. I knew they were there. On some nights, arriving on the At every opportunity, I continued working on the book, 11:00 o’clock train at Frijoles Station on the way to Barro sometimes for months at a time, sometimes in fits and starts Colorado Island, I could hear the last few squeaks and clicks as time allowed. I spent many fulfilling hours in libraries, of the small treefrogs along the shore, or the last plaintive not only at the University of Connecticut, but also at the whines of túngara frogs finishing up a night of calling. Smithsonian Tropical Research Institute during summer I had come to Panama in 1976 as a postdoctoral fellow trips for field research. Over the years, much of the technol- of the Smithsonian Tropical Research Institute. Fresh out of ogy associated with book writing has changed. The initial graduate school, I had little experience in the tropics, but I version of the first chapter I wrote was composed on a knew of the great diversity of frogs to be found there, some- portable electric typewriter, and from there I graduated to times with more species breeding at a single pond than can an IBM Selectric, a mainframe computer terminal, and a be found in the entire northeastern United States. I am not succession of desktop computers. Frequent trips to the li- sure when I first thought of writing a book on the ecology brary to fill notebooks with handwritten notes have largely and behavior of amphibians, but certainly ideas began to been replaced by access to electronic journals and the print- germinate as I took those long train rides along the Panama ing of thousands of pages of recent journal articles. Figures Canal and my many boat rides across Gatun Lake to the drawn by hand with a lettering set have given way to com- jungle island of Barro Colorado. puterized graphics, while the tedious developing of black I actually began writing this book in the early 1980s, so and white photographs in the darkroom has been replaced to say it has had a long gestation period would be a gross by digitally scanned images.
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