SNAKES SNAKES Ecology and Conservation EDITED BY STEPHEN J. MULLIN DEPARTMENT OF BIOLOGICAL SCIENCES EASTERN ILLINOIS UNIVERSITY RICHARD A. SEIGEL DEPARTMENT OF BIOLOGICAL SCIENCES TOWSON UNIVERSITY COMSTOCK PUBLISHING ASSOCIATES A DIVISION OF CORNELL UNIVERSITY PRESS ITHACA AND LONDON Copyright © 2009 by Cornell University All rights reserved. Except for brief quotations in a review, this book, or parts thereof, must not be reproduced in any form without permission in writing from the publisher. For informa- tion, address Cornell University Press, Sage House, 512 East State Street, Ithaca, New York 14850. First published 2009 by Cornell University Press Printed in the United States of America Library of Congress Cataloging-in-Publication Data Snakes : ecology and conservation / edited by Stephen J. Mullin and Richard A. Seigel. p. cm. Includes bibliographical references and index. ISBN 978-0-8014-4565-1 (cloth : alk. paper) 1. Snakes—Ecology. 2. Snakes—Conservation. I. Mullin, Stephen J., 1967– II. Seigel, Richard A. III. Title. QL666.O6S655 2009 597.96'17—dc22 2008046823 Cornell University Press strives to use environmentally responsible suppliers and materials to the fullest extent possible in the publish- ing of its books. Such materials include vegetable-based, low-VOC inks and acid-free papers that are recycled, totally chlorine-free, or partly composed of nonwood fi bers. For further information, visit our website at www.cornellpress.cornell.edu. Cloth printing 10 9 8 7 6 5 4 3 2 1 S. J. M.: In memory of Francis Joseph Mullin, PhD (1906–1997), professor of anatomy and physiology at the University of Chicago, and in memory of his son, my father, Michael Mahlon Mullin, PhD (1937–2000), professor of oceanography at Scripps Institution of Oceanography, for sharing innumerable cultural and educational opportunities with me. R. A. S.: To my parents (Harald and Harriet Seigel) for passing on their love of learning to me, and to Nadia and Ben Seigel for all their love and support. A special dedication to James D. Anderson, whose enthusiasm and love of herpetofauna got all of this started for me. Although his career was cut tragically short, his memory lives on in his students, of which I am proud to have been one. Contents Preface ix Acknowledgments xi List of Contributors xiii Introduction: Opening Doors for Snake Conservation Stephen J. Mullin and Richard A. Seigel 1 Chapter 1. Innovative Methods for Studies of Snake Ecology and Conservation Michael E. Dorcas and John D. Willson 5 Chapter 2. Molecular Phylogeography of Snakes Frank T. Burbrink and Todd A. Castoe 38 Chapter 3. Population and Conservation Genetics Richard B. King 78 Chapter 4. Modeling Snake Distribution and Habitat Christopher L. Jenkins, Charles R. Peterson, and Bruce A. Kingsbury 123 Chapter 5. Linking Behavioral Ecology to Conservation Objectives Patrick J. Weatherhead and Thomas Madsen 149 viii Contents Chapter 6. Reproductive Biology, Population Viability, and Options for Field Management Richard Shine and Xavier Bonnet 172 Chapter 7. Conservation Strategies: Captive Rearing, Translocation and Repatriation Bruce A. Kingsbury and Omar Attum 201 Chapter 8. Habitat Manipulation as a Viable Conservation Strategy Kevin T. Shoemaker, Glenn Johnson, and Kent A. Prior 221 Chapter 9. Snakes as Indicators and Monitors of Ecosystem Properties Steven J. Beaupre and Lara E. Douglas 244 Chapter 10. Combating Ophiophobia: Origins, Treatment, Education, and Conservation Tools Gordon M. Burghardt, James B. Murphy, David Chiszar, and Michael Hutchins 262 Chapter 11. Snake Conservation, Present and Future Richard A. Seigel and Stephen J. Mullin 281 References 291 Taxonomic Index 357 Subject Index 362 Preface This book follows in the footsteps of two previous efforts—Snakes: Ecol- ogy and Evolutionary Biology (1987) and Snakes: Ecology and Behavior (1993)—to provide established and new researchers with a current synopsis of snake ecology. In the preface to each of these earlier works, one of us (R. A. S.) admitted that he had erred in assuming that another “Biology of the Serpentes” book was not worth tackling. And after the fi rst two books, we thought that perhaps yet another book was not needed—we were wrong again. Because our understanding of snake ecology continues to evolve, this fi eld of study provides a seemingly inexhaustible source of research topics to pursue. Furthermore, even more time has now elapsed between this book and its predecessor than between the publications of the fi rst and second books. As such, the need to enlighten our audience about recent advances in method- ology and analysis is obvious. Like the two previous volumes, we developed the concept for this book with three goals in mind: (1) to summarize what is known about the major aspects of snake ecology and conservation, (2) to provide a compilation of the primary literature on this topic that is equally valuable to experienced and developing researchers, and (3) to stimulate new and innovative research on snakes by drawing attention to those areas in which there is a paucity of effort. Given the ever-increasing number of quantifi ed declines in both popula- tion size and species diversity among a variety of taxa, this book has an urgent fourth purpose that almost overshadows the previous three—to pro- vide an awareness of the threats to snake populations and examine the strat- egies available to protect these unique organisms from further population declines or extinction. Indeed, if the reader is familiar with the contents of x Preface the previous snake ecology books, you are already aware that conservation is a topic that carries over from both of them. It is clear to us that the expo- nential growth of the world human population has already exacted a toll, both directly and indirectly, on snake populations. Furthermore, because of their typical role in most trophic webs, it is not a great leap for us to suggest that the health of snake populations is indicative of overall environmental health—in much the same way that amphibians, over the past two decades, have been viewed as the canaries in the environmental coal mine. Other signifi cant events that have occurred since the publication of the second Snakes book include the second through fourth meetings of the Snake Ecology Group, a loosely organized collection of biologists who are united in their enthusiasm for snakes. The attendance and level of participation have increased steadily with each successive conference, and we have ob- served that they are especially conducive to promoting collaborative efforts among several, sometimes disparate, subdisciplines. It is from the presenters at the 2004 meeting that we solicited many of the contributions to this book. Because the fi eld of snake ecology has continued to evolve, it should come as no surprise that the authors of these chapters include many individuals who did not contribute to either of the earlier Snakes books. We encouraged these authors to interact frequently when writing their chapters and to cross- reference one another’s work. As was the case for the two previous books, our primary audience is the professional scientist; we are hopeful that curatorial staff in zoological parks and nongame wildlife managers will also fi nd this information of in- terest. When the previous volumes were published, one of us (S. J. M.) was a student who was further encouraged by them; similarly, we trust this book will stimulate creative research and be an invaluable reference for today’s developing snake ecologists. If nothing else, we hope that our efforts will continue to foster both interest in and scholarship about snake populations with objectives that include their conservation. Stephen J. Mullin Richard A. Seigel Acknowledgments Even though its meetings are irregular, whenever the Snake Ecology Group gets together, one recurring theme is that studying the natural his- tory of snakes is really fun but also potentially challenging because funding is scarce. So, it is only natural for us not only to recognize the excellent work of our authors (and their patience with our requesting multiple revi- sions) but also to acknowledge support provided to all snake ecologists, es- pecially from the ever-decreasing pool of funding agencies that still support research in basic natural history. We also thank all the snake researchers whose work provided the foundations for many of the ideas presented in these chapters. We are grateful for the rewarding interactions with our team at Cornell University Press: Candace Akins, Scott Levine, Heidi Lovette, Susan Specter, and Emily Zoss. In addition to our own internal reviewing process, sev- eral colleagues provided critical feedback at various stages of this project, including G. Blouin-Demers, G. Brown, C. Dodd, H. Greene, J. Mitchell, C. Phillips, H. Reinert, G. Rodda, and J. Rodríguez-Robles. S. J. M. thanks the administration and staff of Department of Biological Sciences at Eastern Illinois University (EIU) for support of this project, and he thanks the students in the EIU Herpetology Lab for their feedback and tolerance of his extended spells of absent-mindedness during its completion. Portions of this book were completed while S. J. M. was on a sabbatical leave granted by Mary Anne Hanner, dean of the College of Sciences, EIU. Previous guidance from mentors during his training (R. Cooper, H. Greene, W. Gutzke, and H. Mushinsky) is also appreciated. R. A. S. thanks Towson xii Acknowledgments University for funding and logistical support during the writing of this book, with special thanks to Dean Intemann, Dean David Vanko, and Pro- vost James Clements. Support for R. A. S. was also provided by the Dyna- mac Corporation, with special thanks to Ross Hinkle for this long-term support. Contributors Omar Attum, Department of Biology, Indiana University Southeast Steven J. Beaupre, Department of Biological Sciences, University of Arkansas Xavier Bonnet, Centre d’Etudes Biologiques de Chizé, Centre National de la Re- cherche Scientifi que (France) Frank T.