Table Of ContentCOMPARATIVE
ANIMAL BEILWIOR
Donald A. Dewsbury
Professor of Psychology
University of Florida
McGRAW-HILL BOOK COMPANY
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COMPARATIVE ANIMAL BEHAVIOR
Copyright " 1978 by McGraw-Hill, Inc. All rights reserved. Printed in the United States
of America. 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 written permission of the publisher.
I234567890FGFG783210987
This book was set in Times Roman by Black Dot, Inc. The editors were Richard R.
Wright and Susan Gamer: the cover was designed by Scott Chelius; the production
supervisor was Dominick Petrellese. The drawings were done by J & R Services, Inc.
Fairfield Graphics was printer and binder.
Library of Congress Cataloging in Publication Data
Dewsbury, Donald A date
Comparative animal behavior.
Bibliography: p.
Includes index.
1. Animals, Habits and behavior of. 2. Psychology,
Comparative. I. Title.
QL751.D452 591.5 77-8885
ISBN 0-07-016673-0
Contents
Preface xi
PART ONE
INTRODUCTION TO ANIMAL BEHAVIOR
Chapter One
The Study of Animal Behavior 3
THE SCOPE OF THE STUDY OF ANIMAL BEHAVIOR 3
EARLY HISTORY 8
SUMMARY 11
Chapter Two
Animal Behavior: An Interaction of Disciplines 12
CLASSICAL ETHOLOGY AND ANIMAL PSYCHOLOGY 12
ETHOLOGY 14
ANIMAL PSYCHOLOGY 22
CONTEMPORARY ETHOLOGY AND ANIMAL PSYCHOLOGY:
A RAPPROCHEMENT 25
vi CONTENTS
CONTRIBUTIONS OF OTHER DISCIPLINES 27
ANIMAL BEHAVIOR TODAY 28
SUMMARY 28
Chapter Three
Animal Species: Their Nature, Classification, and
Evolution 29
ANIMAL SPECIES 29
TAXONOMY 33
EVOLUTION AS HISTORY 34
EVOLUTION AS A PROCESS 36
SUMMARY 46
PART TWO
PATTERNS OF BEHAVIOR
Chapter Four
Individual Behavioral Patterns 49
LOCOMOTION 50
INGESTION 51
THERMOREGULATION 54
SEEKING OF SHELTER 54
AVOIDANCE OF PREDATORS 55
SLEEP 58
BODY MAINTENANCE 59
ELIMINATION 59
EXPLORATION 60
PLAY 63
USE OF TOOLS 64
BIOLOGICAL RHYTHMS 65
SUMMARY 68
Chapter Five
Reproductive Behavior 69
SEXUAL VERSUS ASEXUAL REPRODUCTION 69
BEHAVIORAL PATTERNS ASSOCIATED WITH SEXUAL
REPRODUCTION 70
THREE REPRODUCTIVE CYCLES 83
SUMMARY 88
Chapter Six
Social Organization and Social Behavior 90
SOCIAL ORGANIZATION 91
INDIVIDUAL SOCIAL INTERACTIONS 99
SUMMARY 107
CONTENTS vil
PART THREE
DEVELOPMENT OF BEHAVIOR
Chapter Seven
Behavior Genetics 111
IS OBSERVED BEHAVIOR AFFECTED BY VARIATIONS IN GENOTYPE? 112
HOW MANY GENES ARE OPERATING? 116
WHAT PROPORTION OF OBSERVED VARIABILITY IS ATTRIBUTABLE,
RESPECTIVELY, TO THE GENES AND TO THE ENVIRONMENT? 120
WHAT DEVELOPMENTAL AND PHYSIOLOGICAL PROCESSES
INTERVENE BETWEEN THE GENES AND THEIR EXPRESSION IN OVERT
BEHAVIOR? 124
WHAT CAN BEHAVIOR-GENETIC ANALYSIS TELL US ABOUT ADAPTIVE
SIGNIFICANCE? 127
HOW DO GENES AND THE ENVIRONMENT INTERACT TO PRODUCE
BEHAVIOR? 129
THE ROLE OF BEHAVIORAL GENETICS IN ANIMAL BEHAVIOR 131
SUMMARY 131
Chapter Eight
Experience and Behavioral Ontogeny 132
ONTOGENY OF BEHAVIOR 133
EFFECTS OF ENVIRONMENTAL EVENTS ON BEHAVIORAL ONTOGENY 140
SUMMARY 153
Chapter Nine
Nature and Nurture 154
EARLY HISTORY 155
CLASSICAL ETHOLOGICAL VIEWS 156
CRITICISMS OF CLASSICAL ETHOLOGICAL VIEWS 159
RECENT FORMULATIONS 161
A POINT OF VIEW 163
STUDIES IN BEHAVIORAL DEVELOPMENT 164
SUMMARY 170
PART FOUR
MECHANISMS CORRELATED WITH BEHAVIOR 171
Chapter Ten
Sensory-Perceptual Systems and Function 173
METHODS IN THE STUDY OF SENSATION AND PERCEPTION IN
ANIMALS 175
A BRIEF SURVEY OF SOME SENSORY SYSTEMS 179
DEVELOPMENT OF SENSORY-PERCEPTUAL SYSTEMS 188
NAVIGATION IN BIRDS 191
SUMMARY 195
viil CONTENTS
Chapter Eleven
Neural Mechanisms and Behavior 196
NERVOUS SYSTEMS 196
METHODS IN THE STUDY OF NEURAL MECHANISMS 201
NEUROETHOLOGY 203
NEURAL MECHANISMS AND MAMMALIAN BEHAVIOR 207
SUMMARY 217
Chapter Twelve
Hormones and Behavior 218
HORMONES AND THE REGULATION OF HORMONE SECRETION 219
METHODS IN THE STUDY OF HORMONES AND BEHAVIOR 222
MECHANISMS OF HORMONE ACTION 226
EARLY HORMONE EFFECTS 228
COMPLEXITIES AND INTERACTIONS IN HORMONE ACTION 232
HORMONES AS DEPENDENT VARIABLES 235
A BRIEF SURVEY OF HORMONAL EFFECTS ON DIFFERENT BEHAVIOR
PATTERNS 236
PSYCHONEUROENDOCRINE INTEGRATION 240
SUMMARY 243
PART FIVE
EVOLUTION OF BEHAVIOR
Chapter Thirteen
Evolution of Behavior 247
METHODS IN THE STUDY OF THE EVOLUTION OF BEHAVIOR 248
THE CONCEPT OF HOMOLOGY 250
PRIME MOVERS OF BEHAVIORAL EVOLUTION 253
MICROEVOLUTION OF BEHAVIOR 255
SPECIATION AND REPRODUCTIVE ISOLATION 258
RITUALIZATION AND THE EVOLUTION OF DISPLAYS 261
BEHAVIOR AND SYSTEMATICS 265
DOMESTICATION 267
SUMMARY 270
PART SIX
FUNCTIONS OF BEHAVIOR
Chapter Fourteen
Behavior and Adaptive Significance 273
METHODS 273
STUDIES OF ADAPTIVE SIGNIFICANCE 278
CONTENTS ix
MAINTENANCE OF GENETIC VARIATION 286
ORGANISMS AS ADAPTED WHOLES 288
SUMMARY 292
Chapter Fifteen
Sociobiology 294
CORNERSTONES OF SOCIOBIOLOGY 295
REPRODUCTIVE STRATEGIES 297
SOCIOBIOLOGY, BREEDING SYSTEMS, AND SOCIAL ORGANIZATION 306
SUMMARY 316
PART SEVEN
LEARNING: A SYNTHESIS
Chapter Sixteen
Learning: Description, Development, and Mechanisms 319
DEFINITION OF LEARNING 319
FORMS OF LEARNING 321
DEVELOPMENT OF LEARNING 331
MECHANISMS AND LEARNING 337
SUMMARY 340
Chapter Seventeen
Learning: Evolution and Adaptive Significance 341
EVOLUTION OF LEARNING ABILITY 341
ADAPTIVE SIGNIFICANCE OF LEARNING 356
SUMMARY 363
References 365
Indexes 417
Name Index
Species Index
Subject Index
Preface
This book is intended for use with undergraduates taking their first course in
the area of comparative psychology and animal behavior. It is written by a
comparative psychologist, with a certain emphasis on material from that
discipline. However, the area of animal behavior is so truly interdisciplinary
that any contemporary presentation must represent some kind of synthesis of
material from different related disciplines. Therefore, the book also draws
heavily on material from related disciplines. It is hoped that the book may also
generate some interest in those disciplines.
This book is short, compared with others covering similar material. That is
deliberate: the goal was to create a relatively short book that would provide a
solid core for a course in animal behavior, leaving the instructor free to
supplement this basic core with related readings which would emphasize
material of his or her choosing. Thus, the student would be able to profit from
the instructor's expertise in a particular subarea, and, at the same time, to view
the area of animal behavior in a somewhat broader contemporary perspective.
The basis for the organization of the book lies in the approach to the study
of animal behavior most clearly delineated by Niko Tinbergen. The first task of
the animal behaviorist is to observe and describe behavior. However, observa-
tion and description are not ends in themselves. Rather, they represent a way to
provide a foundation for asking four classes of questions—questions about
xi
xii PREFACE
development, mechanism, evolutionary history, and function. Part One pro-
vides introductory material related to the study of animal behavior. Part Two
comprises three chapters in which behavior is described. The goal is to
familiarize the student with a broad range of behavioral patterns, many of
which will be analyzed further in later sections. Parts Three, Four, Five, and
Six deal with the four classes of questions to be asked of animal behavior—
questions about development, mechanism, evolutionary history, and adaptive
significance. The book concludes with a section on learning (Part Seven). The
organization of Part Seven is parallel to, and restates, that of Parts One through
Six, as it covers description, development, mechanisms, evolutionary history,
and adaptive significance.
I have been fortunate to have had many teachers and colleagues who have
influenced my approach to animal behavior. While I cannot name them all, I
thank them all. The manuscript was typed by Juanita Brown, Donna Gillis, and
Martha Hettrick. Finally, I wish to thank Joyce, Bryan, and Laura for their
tolerance as they discovered that what they may have heard about the
difficulties of having a budding author in the family is all true.
Donald A. Dews bury
