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SNAKES
SNAKES
Ecology and Conservation

E DI TE D 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 fibers. 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 first 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
field 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 first 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 quantified 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 significant 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 field 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 find 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 Scientifique (France)

Frank T. Burbrink, Department of Biology, College of Staten Island–The City Uni-

versity of New York

Gordon M. Burghardt, Departments of Psychology and Ecology & Evolutionary

Biology, University of Tennessee

Todd A. Castoe, Department of Biochemistry & Molecular Genetics, University of

Colorado–School of Medicine

David Chiszar, Department of Psychology, University of Colorado

Michael E. Dorcas, Department of Biology, Davidson College

Lara E. Douglas, Department of Biological Sciences, University of Arkansas

Christopher L. Jenkins, Project Orianne, Ltd.

Glenn Johnson, Department of Biology, State University of New York at Potsdam

Michael Hutchins, The Wildlife Society

Richard B. King, Department of Biological Sciences, Northern Illinois University

Bruce A. Kingsbury, Department of Biology, Indiana University–Purdue University

Fort Wayne
xiv Contributors

Thomas Madsen, School of Biological Sciences, University of Wollongong (Aus-

tralia)

Stephen J. Mullin, Department of Biological Sciences, Eastern Illinois University

James B. Murphy, National Zoological Park

Charles R. Peterson, Department of Biological Sciences, Idaho State University

Kent A. Prior, Critical Habitat, Parks Canada

Richard A. Seigel, Department of Biological Sciences, Towson University

Richard Shine, School of Biological Sciences, University of Sydney (Australia)

Kevin T. Shoemaker, Department of Environmental and Forest Biology, College of

Environmental Science and Forestry–State University of New York

Patrick J. Weatherhead, Program in Ecology & Evolutionary Biology, University

of Illinois

John D. Willson, Savannah River Ecology Laboratory, University of Georgia

SNAKES
 Introduction

Opening Doors for Snake

Conservation

STEPHEN J. MULLIN AND RICHARD A. SEIGEL

An unfortunate certainty associated with the ever-growing human popu-

lation is the loss or alteration of habitat. Coupled with this population in-
crease, technological advances have allowed humans to become more mobile,
and with that mobility comes the increased likelihood that other organisms
will—intentionally or not—move with them. These are just a few of the
reasons why many species of nonhuman organisms are experiencing popu-
lation declines. Although many people are willing to extend some effort for
conservation when endearing animals like pandas or parrots are concerned,
the sympathy extended to the marvelous variety of snake species is rather
limited. This book provides an examination of current research concerning
the ecology of snakes, with an emphasis on how this research has been, or
has the potential to be, applied to their conservation.
Snakes have intrigued humans for centuries, and were incorporated into
several mythologies (e.g., the staff of Aesculapius) and cultures (e.g., Irula
snake-catchers; Whitaker 1989). Among the biologists who study snakes,
there is little question of their fascination about the natural history of
snakes. In spite of a limbless ectothermic body, snake species have radiated
to inhabit all of the Earth biomes except the polar regions— even then, spe-
cies can be found within the Artic circle (e.g., Vipera berus; Carlsson and
Tegelström 2002). The variety of locomotory modes observed in snakes has
garnered much interest (see Gans 1986 and references therein), perhaps ex-
ceeded only by that allocated to snake size–prey size relationships (reviewed
in Arnold 1993). There is also considerable enthusiasm for snakes in a rap-
idly growing and dedicated sector of the commercial pet trade.
2 S. J. Mullin and R. A. Seigel

Sadly, the considerable amount of effort by researchers and enthusiasts

has not translated into public support for snakes. Declines in the sizes of
snake populations do not receive the same level of attention as has recently
been the case for sea turtles (Meylan and Ehrenfeld 2000) or any number
of amphibian species (Miller 2000; Norris 2007). The same enthusiasm for
snakes observed among commercial breeders might be exacting a negative,
but poorly quantified, impact on wild populations (Nilson et al. 1990; Schlaep-
fer et al. 2005). Other human activities are known sources of declines in
wild snake populations (Gibbons et al. 2000), even among venomous spe-
cies (Whitaker and Shine 2000). In the United States, the continued sanc-
tioning of rattlesnake round-ups clearly does not provide any benefits for
the populations of these species (mostly Crotalus adamanteus, C. atrox, and
C. horridus; Fitzgerald and Painter 2000). The troubling nature of this treat-
ment of snakes is compounded by the fact that many of these species repre-
sent the highest levels in their respective trophic webs. As such, continued
declines in snake populations are likely to leave their prey populations (sev-
eral of which are commonly construed as pests) unchecked.
The field of snake ecology has advanced considerably over the last 15
years— conceptual frameworks have been revised in light of new findings,
and improvements in technology have afforded opportunities for new av-
enues of research. The contributors to this book represent a healthy mix
of the seasoned developers of some of these frameworks and techniques,
and the up-and-coming pioneers who have built on these advances to lead
conservation efforts in new directions. In addition to describing some of the
challenges associated with studying snake ecology in Chapter 1, Michael
Dorcas and J.D. Willson discuss several of the recent applications of mark-
ing and modeling snake populations. In Chapter 2, Frank Burbrink and
Todd Castoe not only describe the latest and most appropriate techniques
used in phylogeographic studies, but also tackle the monumental task of
summarizing the recently published research on snake phylogeography. In
Chapter 3, Richard King summarizes the latest work in population genet-
ics and illustrates the processes that generate population structure on fine
geographic and temporal scales. In Chapter 4, Christopher Jenkins, Charles
Peterson, and Bruce Kingsbury couple their expertise with geographical infor-
mation systems and spatial modeling to answer questions associated with
the ecology of snakes at the landscape level.
The next couple of chapters encompass our attempt to update reviews
of areas within snake ecology that have received a fair amount of attention
over the past two decades. In Chapter 5, Patrick Weatherhead and Thomas
Madsen summarize the central concepts in behavioral ecology, with partic-
ular emphasis on thermal ecology and predator-prey interactions. And be-
cause successful reproduction is critical to population viability, we asked
Richard Shine and Xavier Bonnet to interpret the latest research examining
snake reproductive biology (Chapter 6). In spite of these authors’ efforts
 Introduction 3

to combine benchmark studies in these areas with the volume of recent liter-
ature, we are still left with the impression that there is much to learn about
the behavioral and reproductive ecology of snakes, particularly as it per-
tains to their conservation.
We have also included contributions designed to address a few relatively
new, and sometimes controversial, fields that focus specifically on the im-
portance of conserving snake populations in the field. In Chapter 7, Bruce
Kingsbury and Omar Attum discuss the efficacy of management strategies
such as repatriation, translocation, and captive propagation. In Chapter 8,
Kevin Shoemaker, Glenn Johnson, and Kent Prior describe how various
techniques of habitat manipulation can be used to promote the stability of
snake populations or minimize the impacts of human alteration of habitat.
The impacts of snakes in various ecosystems are further illustrated by Ste-
ven Beaupre and Lara Douglas, who describe in Chapter 9 the methodol-
ogy associated with using snakes as biological monitors of environmental
quality.
An enduring mystery to most snake biologists is that the curiosity aroused
in the general public by various aspects of snake biology does not also gen-
erate sympathy for the plight of many of these species. It is for this, and
other reasons, that we have asked Gordon Burghardt, James Murphy, David
Chiszar, and Michael Hutchins to contribute Chapter 10, which examines
human perceptions of, and interactions with, snakes in natural and edu-
cational settings and what can be done to improve the image that snakes
have with the general public. Although the emphasis on conservation might
be perceived as being greater in this chapter, we hope that this theme can
be easily detected in all the contributions to this book.
We expect that this book will be of interest to ecologists, conservation
biologists, and curatorial staff at zoological parks and to be of particular
value to herpetologists and wildlife and resource managers. We especially
dedicate this book to new workers in the field, and we hope that our audi-
ence will share our enthusiasm for snakes and the ecological insights that
have been generated by studying them. Given the amount of information
that is yet to be discovered, we are confident that this book will motivate
future generations of researchers to pursue additional avenues of research
as well as encourage them to advocate the conservation of snakes.
Readers familiar with the first two volumes in this series of books on
snakes might find this one to be lacking in the number of tables that sum-
marize data from the primary literature. Our explanation is that in this
book, to a certain extent, we are navigating in uncharted waters with the
coverage of conservation measures that are specific to snakes. Simply put,
studies addressing the conservation of snakes are relatively few in number,
and many conservation tools that have been applied to other taxa remain to
be tested with snake species. The advances in molecular techniques used to
better understand evolutionary relationships among snake species mandated
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