Evolutionary Behavioral Ecology

660 Pages·2010·4.769 MB·English

by  David Westneat, Charles Fox

EVOLUTIONARY BEHAVIORAL ECOLOGY
Edited by
David F. Westneat
Charles W. Fox
EVOLUTIONARY BEHAVIORAL ECOLOGY
EVOLUTIONARY BEHAVIORAL ECOLOGY
Edited by
David F. Westneat
Charles W. Fox
1
2010
3
Oxford University Press, Inc., publishes works that further
Oxford University’s objective of excellence
in research, scholarship, and education.
Oxford New York
Auckland Cape Town Dar es Salaam Hong Kong Karachi
Kuala Lumpur Madrid Melbourne Mexico City Nairobi
New Delhi Shanghai Taipei Toronto
With offi ces in
Argentina Austria Brazil Chile Czech Republic France Greece
Guatemala Hungary Italy Japan Poland Portugal Singapore
South Korea Switzerland Thailand Turkey Ukraine Vietnam
Copyright © 2010 by Oxford University Press, Inc.
Published by Oxford University Press, Inc.
198 Madison Avenue, New York, New York 10016
www.oup.com
Oxford is a registered trademark of Oxford University Press.
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
electronic, mechanical, photocopying, recording, or otherwise,
without the prior permission of Oxford University Press.
Library of Congress Cataloging-in-Publication Data
Evolutionary behavioral ecology / edited by David F. Westneat, Charles W. Fox.
p. cm.
Includes bibliographical references and index.
ISBN 978-0-19-533193-6; 978-0-19-533192-9 (pbk)
1. Animal behavior—Evolution. 2. Animal ecology.
I. Westneat, David F. II. Fox, Charles W.
QL751.E87 2010
591.5—dc22 2009031325
9 8 7 6 5 4 3 2 1
Printed in the United States of America
on acid-free paper
Preface
An understanding of the natural world and what’s in it is a
source of not only a great curiosity but great fulfi llment.
—Sir David Attenborough, September 2004, during a
break from fi lming bolas spiders at the University of
Kentucky’s Spindletop Farm
The diversity of ways in which organisms inter- the latest in 1997, the Krebs and Davies volumes
act with their environment, including members brought both conceptual clarity and empirical
of their own species, other species, and the abiotic excitement to the fi eld. Probably every practic-
world, is astounding. The fi eld of behavioral ecol- ing behavioral ecologist has gained insights from
ogy explores the role of behavior in these interac- these volumes. Although these editions still make
tions. Behavioral ecologists are typically interested for compelling reading and provide excellent over-
in a functional understanding of behavior: why views of the logic and methods of the fi eld, they
do animals behave the way they do, and make no longer refl ect the full scope of the fi eld: some of
the decisions that they make? How are animal the material is dated, and each edition intentionally
decisions and behaviors shaped by evolution in included an eclectic set of topics not broad enough
response to ecological and/or social conditions that in scope to form the foundation for a comprehen-
they experience? Such questions have turned out sive, semester-long graduate class.
to be surprisingly complicated, requiring a diver- In Evolutionary Behavioral Ecology we have
sity of approaches, from mathematical modeling to attempted to present the core concepts and recent
biogeographic analyses to fi eld manipulative stud- elaborations upon them that defi ne the fi eld today—
ies, and integration among fi elds of study, includ- a daunting task because the fi eld has grown sub-
ing psychology, evolutionary biology, physiology, stantially since Krebs and Davies’ fi rst publication.
molecular biology, and even economics, among Their volumes developed in detail select topics that
others. The insights gained by behavioral ecologists highlighted major conceptual developments in the
have generated considerable controversy over the fi eld, introducing these to new behavioral ecologists
years, spurring the fi eld to move in new directions. and energizing the veterans. However, the fi eld is too
Two of the most deservedly famous individu- rich and deep to simultaneously introduce begin-
als in behavioral ecology are John Krebs and Nick ning students to the diversity of ideas that make
Davies. Though the study of behavioral ecology did up modern behavioral ecology, and present each of
not start with them, its defi nition as a fi eld of study those fi elds with the thoroughness of a Krebs and
was in large part due to their immensely infl uen- Davies chapter. To accomplish our goal of making
tial book, Behavioural Ecology: An Evolutionary a volume accessible to students, we asked authors
Approach, published in 1978. That single book to focus on core concepts, presented rigorously, and
pulled together the variety of behaviors and ideas not write encyclopedic reviews. We wanted authors
that had been the focus of researchers interested to target their chapters directly at fi rst- or second-
in functional explanations. Through four editions, year graduate students and lead them through the
vi Preface
main ideas that have shaped the fi eld over the past themes as well as the personalities involved in its
40 (or more) years, highlighting some of the major development. We thus start with a chapter on the
controversies that excite us today. Our authors history of behavioral ecology, writ large (chapter 1),
were thus challenged to sacrifi ce detail and instead before delving into more specifi c subjects. The rest
focus on major ideas that have driven the fi eld for- of sections I and II provide students with an intro-
ward. We hope students fi nd these brief introduc- duction to the tools and broad conceptual ideas
tions stimulating and that they pursue additional that underpin the entire fi eld, regardless of specifi c
readings, such as those listed in “Suggestions for subject. Together, the two sections are deliberately
Further Reading” sections, to explore ideas in more linked, albeit implicitly and somewhat loosely, to
detail. Most important, we hope that readers of this Tinbergen’s (1963) four questions: mechanism
book—students of behavioral ecology—will take (chapters 9 and 10), development (chapters 5 and
away a sense of the vitality of a fi eld full of bril- 6), evolution (chapter 7), and function (chapters 2
liant insights but also brimming with unanswered and 8).
questions that students themselves can contribute In sections III, IV, and V we march through the
to answering. traditional topics in behavioral ecology: fi nding
Unlike many books targeted to students and the food and avoiding predators (section III), social
classroom, we chose to edit a collection of chap- behavior (section IV), and reproductive behavior
ters written by others rather than to write a book (section V). We have sequenced chapters to provide
ourselves. A multiauthored book has strengths and broad, overarching concepts early and elaborations
weaknesses. One strength is that these chapters are on those themes later in each section. Section VI
written by experts in their respective areas, and so is called “Extensions,” but it could also be called
each represents a mature perspective on the impor- “Cross-Cutting” or “Emerging Frontiers.” Here
tant topics. A potential weakness of multiauthored authors take key ideas from behavioral ecology and
books is an inconsistency of style, level of mate- apply them to subjects such as speciation (chapter
rial, and breadth of focus among chapters. We 27), conservation biology (chapter 29), and human
encouraged authors to follow a similar outline for behavior (chapter 31), or they present new concep-
chapters, and we aggressively edited those chapters tual views (e.g., chapter 30 on personality) or tech-
to provide some consistency and to highlight con- niques (chapter 28 on genomics) that seem likely
nections among chapters, but variation in style and to provide behavioral ecologists with new ways of
level of presentation still exists. Some of these dif- elaborating on functional approaches.
ferences, such as how much mathematical modeling Of necessity, we could not cover all subfi elds of
is included, we think are a strength of an edited behavioral ecology and, for most areas, we have
volume because this diversity refl ects real variation touched only briefl y on the diversity of exciting
within the fi eld, both among subjects and among questions being asked. Moreover, we have asked
researchers within subjects: some researchers do authors to be sparing in their use of citations. We
quite well without using mathematical modeling did this to encourage a focus on concepts and key
and others depend on it, yet everyone gains under- developments and to limit the degree to which
standing from it. Chapters will not all be equally chapters became encyclopedic reviews. We regret
accessible to the intended audience, and some if some subjects and key papers thus appear to be
authors have perspectives on their topic that differ ignored, and we apologize to those whose favorite
from even the majority of their colleagues (though example, case study, or specifi c citation was left out
we’ve strived for balanced perspectives as much of the book.
as possible). Nevertheless, we think both of these There are a few unique elements of this book
problems are potentially benefi ts. Students grow that reaffi rm to us the value of having a new sum-
as professionals by being confronted with topics in mary of our fi eld every 10–12 years. Just as each
which there is confusion and disagreement among edition of Krebs and Davies highlighted new ideas
professionals. Often such areas of disarray are fer- and approaches that were changing the future of
tile ground from which new careers can sprout. behavioral ecology, the emergent picture of behav-
Behavioral ecology arose from scientists mixing ioral ecology from our book is one of increasing
questions and approaches among disciplines, and to sophistication of both questions and techniques. By
understand the state of the fi eld today it is instruc- far the most compelling advances in the fi eld have
tive to students to learn a bit about the historical been due to the increased use of genetic tools used
Preface vii
to assess relatedness both within populations (e.g., Anders Berglund, Andrew Bourke, Tim Caro, Will
kinship) and among taxa (as a basis for construct- Creswell, Jonathon Crystal, Reuven Dukas, Daniel
ing phylogenies). These approaches have become Funk, Carl Gerhardt, Greg Grether, Paul Harvey,
so well ensconced in the fi eld they are hardly men- Ken Haynes, Natalie Hempel de Ibarra, Geoff Hill,
tioned explicitly, but their impact is broad. More Jerry Husak, Adam Jones, Laurent Keller, Darrell
compelling is the increased use of molecular tools, Kemp, Dov Lank, Kate Lessells, Wayne Linklater,
as described in chapters 5 and 28, for understand- Jeffrey Lucas, Joel McGlothlin, Molly Morris,
ing the genetic basis of behaviors of ecological rel- Bryan Neff, Geoff Parker, Tomasso Pizzari, Denis
evance, and the insights these provide regarding Reale, David Reznick, GraemeRuxton, Mike Ryan,
function and evolution of behavior. Finally, new William Searcy, Maria Servedio, Mike Sharkey,
advances in quantitative genetics have great poten- Ulrich Steiner, William Wagner, Jonathan Wright,
tial to alter conceptual approaches and empirical and Marlene Zuk. We also thank Dov Lank and
research on a wide array of behaviors. General the- his graduate class in behavioral ecology at Simon
ory on plasticity (chapter 6) and on indirect genetic Fraser University for providing extensive feedback
effects (chapter 14) enhances many traditional on many of the chapters. Several members of our
approaches to behavioral ecology and is likely to labs, notably Amanda Ensminger, Damon Orsetti,
infl uence many future studies. Sarah Martin, Bridget Sousa, Patricia Hartman, Ian
Preparing this book involved the assistance of Stewart, and Dan Wetzel, read many of the chapters
many individuals. First and foremost, we thank the at several stages and provided numerous helpful
authors, who have produced stimulating chapters comments. The Department of Entomology at the
and have responded well to the many demands University of Kentucky provided fi nancial support.
we placed on them. All chapters were reviewed Both our research programs have been supported
both internally (by authors of other chapters) and by the National Science Foundation, and that sup-
externally; in addition to the authors who worked port helped us to plan and carry out this project.
as reviewers, and a few anonymous reviewers, we
thank the following people for their helpful com- David F. Westneat
ments: Sigal Balshine, Alex Basolo, Guy Beauchamp, Charles W. Fox
This page intentionally left blank
Contents
Contributors xiii
Section I. Foundations
1. Ingenious Ideas: The History of Behavioral Ecology 3
Tim R. Birkhead and Pat Monaghan
2. Adaptation 16
Charles W. Fox and David F. Westneat
Box 2.1. Optimality Models 26
Steven Hecht Orzack
3. Behavioral Concepts of Selection 32
Barry Sinervo and Ryan Calsbeek
4. What Is Fitness, and How Do We Measure It? 46
John Hunt and David Hodgson
5. The Genetic Basis of Behavior 71
Kerry L. Shaw and Chris Wiley
Box 5.1. A Brief Introduction to Quantitative Genetics 75
Jason B. Wolf and Allen J. Moore
Box 5.2. Diversity of Sex-Determining Mechanisms 81
Daniel A. Warner and Fredric J. Janzen
6. Behavior as Phenotypic Plasticity 90
Cameron K. Ghalambor, Lisa M. Angeloni, and Scott P. Carroll
Box 6.1. Contrasting Quantitative Genetic Models for the Evolution of Plasticity 92
Box 6.2. Contrasting Statistical Methods for Studying Phenotypic Plasticity 96