Animal Behavior 11th Edition John

Alcock
Visit to download the full and correct content document:
https://textbookfull.com/product/animal-behavior-11th-edition-john-alcock/
More products digital (pdf, epub, mobi) instant
download maybe you interests ...

Animal Behavior: An Evolutionary Approach 11th Edition

Dustin R. Rubenstein And John Alcock

https://textbookfull.com/product/animal-behavior-an-evolutionary-
approach-11th-edition-dustin-r-rubenstein-and-john-alcock/

Domestic Animal Behavior for Veterinarians and Animal

Scientists, 6th Edition Katherine A. Houpt

https://textbookfull.com/product/domestic-animal-behavior-for-
veterinarians-and-animal-scientists-6th-edition-katherine-a-
houpt/

Principles of Animal Behavior Fourth Edition Lee Alan

Dugatkin

https://textbookfull.com/product/principles-of-animal-behavior-
fourth-edition-lee-alan-dugatkin/

Animal behavior: concepts, methods, and applications

Second Edition Nordell

https://textbookfull.com/product/animal-behavior-concepts-
methods-and-applications-second-edition-nordell/
Animal Neopragmatism: From Welfare to Rights John
Hadley

https://textbookfull.com/product/animal-neopragmatism-from-
welfare-to-rights-john-hadley/

Comparative Government and Politics 11th Ed John

Mccormick

https://textbookfull.com/product/comparative-government-and-
politics-11th-ed-john-mccormick/

Small Animal Imaging: Self-Assessment Review 1st

Edition John S. Mattoon

https://textbookfull.com/product/small-animal-imaging-self-
assessment-review-1st-edition-john-s-mattoon/

Mathematics Rebooted A Fresh Approach to Understanding

1st Edition Lara Alcock

https://textbookfull.com/product/mathematics-rebooted-a-fresh-
approach-to-understanding-1st-edition-lara-alcock/

Behavior Management: From Theoretical Implications to

Practical Applications 3rd Edition John W. Maag

https://textbookfull.com/product/behavior-management-from-
theoretical-implications-to-practical-applications-3rd-edition-
john-w-maag/
 ...

Animal
Behavior
ELEVENTH EDITION

Dustin R. Rubenstein • John Alcock

Animal Behavior ELEVENTH EDITION

Dustin R. Rubenstein • John Alcock

Columbia University Arizona State University

m SINAUER ASSOCIATES

NEW YORK OXFORD

OXFORD UNIVERSITY PRESS

-w
i publishing worldwide. Oxford is a registered trade mark of Oxford University Press
■
m 198 Madison Avenue, New York, NY 10016, United States of America
Animal
Behavior
ELEVENTH EDITION
Dustin R. Rubenstein • John Alcock
About the Cover
This male peacock spider (Maratus
elqjhans) from Tamworth, NSW,
Australia, unfurls from underneath his
abdomen what looks like the tail of a
male peacock. He also raises his legs as
if to dance, something you can watch
by following the QR code on the back
cover. (Photograph by Michael Doe.)
Animal Behavior, Eleventh Edition
1 Oxford University Press is a department of the University of Oxford. It furthers
the University's objective of excellence in research, scholarship, and education by
in the UK and certain other countries.
Published in the United States of America by Oxford University Press
© 2019,2013,2009,2005,2001,1998,1993,1989,1984,1979,1975 Oxford University Press
Sinauer Associates is an imprint of Oxford University Press.
For titles covered by Section 112 of the US Higher Education Opportunity Act,
please visit www.oup.com/us/he for the latest information about pricing and
alternate formats.
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, without the prior
permission in writing of Oxford University Press, or as expressly permitted by
law, by license, or under terms agreed with the appropriate reproduction rights
organization.
Inquiries concerning reproduction outside the scope of the above should be sent to
the Rights Department, Oxford University Press, at the address above.
You must not circulate this work in any other form and you must impose this same
condition on any acquirer.
Address editorial correspondence to:
Sinauer Associates
23 Plumtree Road
Sunderland, MA 01375 USA
publish@sinauer.com
Address orders, sales, license, permissions, and translation inquiries to:
Oxford University Press USA
2001 Evans Road
Cary, NC 27513 USA
Orders: 1-800-445-9714
Library of Congress Cataloging-in-Publication Data
Names: Alcock, John, 1942- author. | Rubenstein, Dustin R., author.
Title: Animal behavior / Dustin R. Rubenstein, Columbia University,
John Alcock, Arizona State University.
Description: Eleventh edition. | Sunderland, Massachusetts: Oxford
University Press, [2019] | Revised edition of: Animal behavior: an
evolutionary approach / John Alcock. 10th ed. c2013. | Includes
bibliographical references.
Identifiers: LCCN 20180045801 ISBN 9781605355481 (paperback) | ISBN
9781605357706(ebook)
Subjects: LCSH: Animal behavior-Evolution.
Classification: LCCQL751 .A58 2019 | DDC 591.5-dc23
LC record available at https://lccn.loc.gov/2018004580
98765432
Printed in the United States of America
 To my parents,
zoho inspired in me a love of nature
and animal behavior at an early age.
DRR

V*:
 I

Contents in Brief
CHAPTER 1 An Introduction to Animal Behavior 2
CHAPTER 2 The Integrative Study of Behavior 24
CHAPTER 3 The Developmental and Genetic
Bases of Behavior 58
CHAPTER 4 The Neural Basis of Behavior 104
CHAPTER 5 The Physiological Basis of Behavior 144
CHAPTER 6 Avoiding Predators and Finding Food 184
CHAPTER 7 Territoriality and Migration 218
CHAPTER 8 Principles of Communication 256
CHAPTER 9 Reproductive Behavior 302
CHAPTER 10 Mating Systems 352
CHAPTER 11 Parental Care 400
CHAPTER 12 Principles of Social Evolution 444
CHAPTER 13 Social Behavior and Sociality 476
CHAPTER 14 Human Behavior 512
CHAPTER
Art Introduction to Animal Behavior
Natural Selection and the Evolution
of Behavior 5
The Cost-Benefit Approach to
Behavioral Biology 7
The Levels of Analysis 8
Box 1.1 Natural selection and
infanticide in primates 9
The Integrative Study of Animal
Behavior 11
The Integrative Study of Behavior
The Development of Song
Learning 26
Box 2.1 Characterizing sounds
made by animals 27
Intraspecific Variation and
Dialects 28
Social Experience and Song
Development 33
Mechanisms of Song Learning
Box 2.2 Song learning in birds
adoptedby another species 35
The Genetics of Song Learning 36
Control of the Avian Song
System 36
Box 2.3 Proximate mechanisms
underlying song preferences in
females 37
The Evolution of Song Learning
An Evolutionary History of Bird
Song 40
2
Approaches to Studying
Behavior 13
The Adaptive Basis of Behavior:
Mobbing in Gulls 13
Box 1.2 Phylogenies and the
comparative method 14
Box 1.3 The benefit of high nest
density for the arctic skua 18
The Science of Animal Behavior 21
24
Mechanisms of Song Learning and
the Comparative Approach 42
Human versus Avian Vocal
Learning 44
The Adaptive Value of Song
Learning 44
Box 2.4 Why might song learning
make males communicate more
34 effectively with rivals or potential
mates? 45
Adapting to the Local
Environment 46
Recognition: Friends versus
Foes 46
Sexual Selection: Male-Male
Competition 50
Sexual Selection: Female Choice
and Assortative Mating 51
40
The Integrative Study of Bird
Song 56
i
I
VIII CONTENTS
The Developmental and
Genetic Bases of Behavior
Behavior Requires Genes
and the Environment 60
The Interactive Theory
of Development 61
Box 3.1 Behavioral genetics:
Identifying the genetic basis of
differences in behavior 62
Environmental Differences Can
Cause Behavioral Differences 68
Genetic Differences Can Also Cause
Behavioral Differences 69
Box 3.2 Migratory
restlessness 71
Learning and Cognition 74
Learning Requires Both Genes and
Environment 75
Learning in Complex
Environments 76
The Neural Basis of Behavior 104
Responding to Stimuli 106
Complex Responses to Simple
Stimuli 107
How Moths Avoid Bats 110
Box 4.1 Ultrasound detection
in the moth ear 114
Ultrasonic Hearing in
Other Insects 117
Neural Command and Control 118
Decision Making in the Brain
From Ultrasound to Ultraviolet
Radiation 120
Selective Relaying of Sensory
Inputs 124
58
The Adaptive Value of Learning 78
The Evolutionary Development
of Behavior 84
The Evo-Devo Approach to
Understanding Behavior 84
Box 3.3 The genetics of foraging
behavior in honey bees 85
Early Life Developmental
Conditions 89
The Role of the Social
Environment 90
Developmental Homeostasis versus
Developmental Constraint 91
Developmental Switch
Mechanisms 95
Supergenes and Behavioral
Polymorphisms 100
Responding to Relayed
Messages 126
The Proximate Basis of Stimulus
Filtering 129
Box 4.2 Determining how female
parasitoid wasps choose their
singing male bush-cricket
hosts 132
Box 4.3 Cortical magnification
in mammals 135
118 The Evolution of Cognitive
Skills 135
Box 4.4 Do energetic demands
explain why humans have such
large brains? 137
i
 CONTENTS IX

CHAPTER

5 The Physiological Basis of Behavior 144

Endogenous Rhythms and Changing Social Conditions and
Behavioral Priorities 146 Changing Priorities 165

mm Mechanisms of Changing Behavioral

Priorities 146
The Neurobiology of Circadian
Timing 149
Hormonal Mechanisms Underlying
Behavioral Change 166
Organizational versus Activational
Effects of Hormones on Behavior
The Genetics of Circadian and Development 166
Timing 151 Box 5.2 Measuring hormones in
The Physiology of Circadian animals 168
Timing 152 Box 5.3 Do steroid hormones
Seasonal and Annual Cycles modulate male parental behavior
of Behavior 155 in California mice? 171

Cues That Entrain Cycles of Hormones and Reproduction 172

Behavior 156 Testosterone and Reproductive
Predictable Environmental Cues -157 Behavior 174

Box 5.1 Hormonal responses to The Costs of Hormonal

Regulation 176
light in birds 160
Glucocorticoids and Responding to
Unpredictable Environmental
Environmental Change 179
Cues 162

CHAPTER

6 Avoiding Predators and Finding Food 184

Avoiding Predators 186 Finding Food 204
! Social Defenses 186 Optimality Theory and Foraging
r*-- Decisions 205
Box 6.1 Evolutionary game
theory 187 Box 6.3 Territoriality and feeding
I -I ____
behavior in golden-winged
Game Theory and
sunbirds 206
Social Defenses 192
Box 6.4 Optimal foraging by pike
Box 6.2 Game theory and
cichlid fish 208
the selfish herd 192
Criticisms of Optimal Foraging
Blending In 193
Theory 211
Standing Out 197
Landscapes of Fear 213
Optimality Theory and Antipredator
Game Theory and Feeding
Behavior 202
Behavior 214
 X CONTENTS
:

\
Territoriality and Migration 218
Where to Live 220 Box 7.2 Opposite patterns of sex- !
Habitat Selection 220 biased dispersal in mammals
and birds 238
Territoriality and Resource-Holding
Potential 222 Migration 240
i
Box 7.1 How to track migratory The Costs and Benefits of
Migration 243 i
songbirds 228
Why Give Up Quickly When Fighting Box 7.3 Behaviors to reduce
:
for a Territory? 229 the costs of flying during
migration 244
The Dear Enemy Effect 235
Variation in Migratory Behavior 250
To Stay or Go 235 Box 7.4 Migratory pathways of
Dispersal 237 Swainson’s thrush 253 j
!

:;?rzr:r-
Principles of Communication
_______________________________
256 HBSOffIKsssfi's_ 'i

Communication and Box 8.2 Why do female moths

Animal Signals 258 mate with males that produce
Information Use and ultrasonic mimetic signals similar i
Animal Signals 259 to those produced by predatory
bats? 275
The Evolution of Animal
Signals 262 The Function of Animal Signals 278
The Adaptive Function of ?
Preexisting Traits and
the Development of a Strange Display 278
a Strange Display 264 Honest Communication and i
The Panda Principle and Threat Displays 281
!
Preexisting Traits 265 Honest Signaling 284 I
Preexisting Biases and the Evolution Box 8.3 Mechanisms and
of Animal Signals 269 measurement of animal
Box 8.1 Spiders hunting prey coloration 287 :
at night 271 When Multiple Honest Signals Are
Preexisting Traits versus Better Than One 288
Preexisting Biases 273 Deceitful Signaling 290
Eavesdropping on Others 295
I

i
:
I)

I
!

I
i

I L

CHAPTER
9 Reproductive Behavior
Sexual Selection and the Evolution
of Sex Differences 305
Sex Differences in Reproductive
Behavior 307
Box 9.1 Are sperm always
cheap? 308
Sex Differences and Parental
Investment 309
A Reversal in Sex Differences 310
Intrasexual Selection and
Competition for Mates 313
Competition and Access to
Mates 314
Coexistence of Conditional
Mating Tactics 316
Coexistence of Alternative
Mating Strategies 320
Sperm Competition 322
CHAPTER
10 Mating Systems 352
Monogamy: A Lack of Multiple
Mating 355
Why Be Monogamous? 355
Monogamy in Species with
Paternal Care 358
Monogamy When Paternal Care
Is Rare 360
Polyandry: Multiple Mating
by Females 362
Monogamous Males and
Polyandrous Females 362
Box 10.1 Sexual parasitism, dwarf
males, and the evolution of
gigolos 364
Polyandry and Indirect Genetic
Benefits 367
Box 10.2 Extra-pair paternity and
good genes in birds 371
CONTENTS XI
302
Mate Guarding and Paternity
Assurance 325
Intersexual Selection and
Mate Choice 328
Female Mate Choice for
Direct Benefits 329
Female Mate Choice for
Indirect Benefits 334
Box 9.2 Sexual selection in
the peacock 337
Runaway versus Chase-away
Sexual Selection 341
Cryptic Female Choice 343
Sexual Conflict 344
The Manipulation of Female
Choice 344
Sexual Arms Races 347
Polyandry and Direct Benefits 377
Polygyny: Multiple Mating by
Males 380
Female Defense Polygyny 381
Resource Defense Polygyny 383
Lek Polygyny 386
Scramble Competition
Polygyny 392
Box 10.3 Lekking females in a sex-
role reversed pipefish 394
Polygynandry and Promiscuity:
Multiple Mating by Both
Sexes 396
Polygynandry 396
Promiscuity 397
XII CONTENTS

CHAPTER

11 Parental Care 400

Offspring Value and Parental
Investment 402
Parental Care Decisions 402
Parental Favoritism in Offspring Care
and Production 407
Parental Favoritism in Humans 409
Family Conflict 411
To Care or Not to Care 414
The Costs and Benefits of
Parental Care 414
Sexual Conflict and Parental Care:
Who Cares? 415
Box 11.1 Why do females provide all
of the care in treehoppers? 418
Why Do Females Care? 419
Why Do Males Care? 421
Box 11.2 Reactions of nest-
defending bluegill males to
potential egg and fry predators
under two conditions 422
Discriminating Parental Care 427
Recognizing One’s Own
Offspring 427
Box 11.3 Why do parents in some
species adopt genetic strangers
of their own species? 430
Interspecific Brood Parasitism 430
Choosing the Correct Host 434
Coevolutionary Arms Races 435
The Evolution of Interspecific Brood
Parasitism 439

Principles of Social Evolution 444

Box 12.1 The major evolutionary
transitions 47
Altruism and the Levels of
Selection 448
Individual versus Group Selection
449
Altruism and the Role of Kin
Selection 450
Box 12.2 Calculating genetic
relatedness 452
Kin Selection and Inclusive Fitness
Theory 453
Challenges to Kin Selection and
Inclusive Fitness Theory 455
Box 12.3 Altruism in
amoebae 455
Haplodiploidy and the Evolution of
Eusociality 458
Testing the Haplodiploidy
Hypothesis 460
Inclusive Fitness and Monogamy in
Eusocial Insects 461
Box 12.4 Division of labor in clonal
trematode flatworms 463
Sterility and Caste
Differentiation 467
Social Conflict in Animal
Societies 470
Reproductive Conflict 470

■A

Social Behavior and Sociality 476
The Evolution of Social
Behavior 478
Forms of Social Behavior 479
Mutual Benefit 479
Box 13.1 How do groups of animals
decide where to go? 480
Box 13.2 Social network
analysis 485
Altruism and Reciprocity 487
Non-cooperative Social Behaviors:
Selfishness and Spite 491
Individual Differences in Social
Behavior 491
Personalities in Social Species 492
The Evolution of Cooperative
Breeding 493
CHAPTER
Human Behavior 512
Communication 514
The Development and Evolutionary
History of Human Speech 515
Box 14.1 Ethical studies of humans
and other animals 517
The Neurophysiology of Speech 519
The Adaptive Value of Speech 521
Reproductive Behavior 523
An Evolutionary Analysis of Human
Mate Choice 524
Mate Choice by Women 524
Box 14.2 Female choice and the
features of dominant versus
attractive men 526
CONTENTS XIII
Reproductive Cooperation and Kin
Selection 494
Reproductive Benefits and
Cooperative Breeding 495
Box 13.3 Mobbing and kinship in
groups of Siberian jays 496
Reproductive Costs and Cooperative
Breeding 500
Reproductive Conflict in
Cooperative Breeders 504
Reproductive Suppression 504
Reproductive Skew, Extra­
pair Paternity, and Social
Structure 507
Box 13.4 Why do males and
females both have elaborate
traits in social species? 508
■■■■■
■■■HI
---------- ----------------------------—
Box 14.3 Human mate choice in an
online world 528
Mate Choice by Men 532
Reproductive and Sexual Conflict in
Humans 535
Extreme Sexual Conflict in Humans:
Polygamy and
Extramarital Affairs 538
Coercive Sex 540
Practical Applications of Behavioral
Theory 543
Evolutionary Medicine 544
The Triumph of an Evolutionary
Analysis of Human Behavior 546
 Preface
For more than 40 years, Animal Behavior has been the leading textbook for intro­
ducing undergraduate students to the topic of animal behavior. John Alcock
authored the first edition of this book in 1975, and after 9 subsequent versions,
this eleventh edition brings on a new lead author, Dustin Rubenstein. The elev­
enth edition maintains its narrative tone as well as its focus on both evolution­
ary and mechanistic approaches to understanding how and why animals as
different as insects and humans behave the way they do. In an effort to keep up
with the rapidly evolving field of animal behavior, this new version also brings
a more integrative approach to studying behavior, emphasizing the growing
body of research linking behavior to the brain, genes, and hormones, as well as
to the surrounding ecological and social environments. Topics like epigenetics
and collective behaviors are highlighted for the first time. Additionally, the
book covers the growing number of comparative phylogenetic studies in ani­
mal behavior that make use of ever-larger molecular phylogenies to generate
and test new ideas in the evolution of animal behaviors. Ultimately, the book
retains its primary goal of giving students a window into the various level
of analysis that researchers use to explain why all living things—including
humans—behave, often in complex ways.

New to the Eleventh Edition

In addition to a new lead author, the eleventh edition features several new
approaches and features that support both student learning and instructor
teaching. After extensive research concluded that most users prefer the classic
organizational structure from early editions, rather than the changes that were
made in the tenth edition, the book returns to its classic organizational struc­
ture, with proximate mechanisms introduced early in the book and before an
extended discussion of the ultimate factors underlying behavior. Yet, each chap­
ter attempts to highlight both proximate and ultimate explanations throughout,
illustrating the integrative nature of the field today. This edition also includes a
refined organizational structure within each chapter that makes material more
accessible to students and instructors alike. Figures and photos throughout the
book have been updated and revised as well, providing a fresher look that,
along with the addition of error bars on most graphical figures, makes it easier
for students to learn how to interpret data. The addition of new pedagogi­
cal features also makes it easier for students to learn complex concepts and
apply behavioral and evolutionary thinking to thought-provoking questions
and problems. Found in every chapter, these new boxes and tables emphasize
hypothesis testing, data interpretation, and problem solving. Finally, since most
students are drawn to animal behavior out of a fascination with something they
have seen on TV or in nature, in each chapter we use QR codes offset in the
margins to give students rapid access to high-definition video and audio clips
of the behaviors discussed in the text.

,
V

PREFACE XV

Focus on Integration
This book provides a comparative and integrative overview of animal behavior,
linking a diversity of behaviors and their adaptive functions to the brain, genes,
and hormones, as well as to the surrounding ecological and social environ­
ments. Just as so many modern studies in animal behavior are taking advan­
tage of new neurobiological or molecular approaches, this book introduces
these and other cutting-edge techniques to its readers, all while maintaining
a focus on the theoretical aspects of the field in an explicit hypothesis-testing
framework. Ultimately, the book highlights both the evolutionary and mecha­
nistic approaches to studying animal behavior, as well as the interdisciplinary
approaches that emphasize the neural, genetic, and physiological mechanisms
underlying adaptive behaviors. Because this new edition of the book is more
integrative than previous versions, we have dropped the book's subtitle, An
Evolutionary Approach, and now simply call it Animal Behavior. The concept of
evolution by natural selection remains our guide as we expand in new and
exciting directions.
In addition to the focus on integrative approaches, this edition features an
update of specific topics such as how new technologies are revolutionizing the
way we study animal movements and the genetic architecture of behavior, as
well as new and expanded coverage of animal social behavior, animal com­
munication, and human behavior, in addition to other topics. New empiri­
cal examples have been added to each chapter, some of which extend classic
examples in new and exciting directions.

New Pedagogical Approach

Built upon the foundation of rich content and discussion topics from previ­
ous editions, we've applied more modem pedagogical tools and added new
features in the eleventh edition. Found in every chapter, these new boxes and
tables emphasize hypothesis testing, data interpretation, and problem solving.
These features are designed to engage students, both inside and outside of the
classroom, and aid instructors in developing their courses and delivering their
lectures. New features include:
• INTEGRATIVE APPROACHES: Focusing attention on cutting-edge
tools in the study of animal behavior, these boxes cover tools for study­
ing birdsong, animal coloration, behavioral genetics, hormones, and
how to conduct ethical studies of humans, among other topics.
• HYPOTHESES TABLES: In brief summaries, alternative or non-
mutually exclusive hypotheses for specific animal behaviors are pre­
sented side-by-side.
• EXPLORING BEHAVIOR BY INTERPRETING DATA: In these boxes,
we explore a concept in data analysis, such as analyzing spectrograms
to show how song learning in birds can be adopted by other species, or
drawing conclusions and generating new hypotheses from empirical
data figures.
XVI PREFACE

• DARWINIAN PUZZLES: Adapted from this popular feature in the

tenth edition, these boxes deal specifically with unresolved "puzzles"
in behavioral research, such as why females in some species have what
looks like a penis but males don't.
In most boxes, students are challenged to "Think Outside the Box" by
answering a series of thought-provoking questions related to the content and
figures inside the box. These have been designed to foster in-class discus­
sion between instructors and students. This feature complements Discussion
Questions, which have now been consolidated online for easier student and
instructor access.

QR Codes/Direct Web Links for Audio and Video Clips

Students taking a course on animal behavior want to see and hear animals
behaving, and in this edition, we've integrated that directly into the text in
an easily accessible manner. Using QR codes (and short URLs), students can
immediately access audio and video directly related to the text in real-time,
without having to wade through a large collection of resources on a companion
website.

ABOUT THE TECHNOLOGY. On many iPhones and some Android phones,

QR codes no longer require a QR reader app. (Any iPhone running iOS 11 or
newer can read QR codes directly.) Simply open the camera, hold the phone
over the QR code, and the audio/video link will appear. Other smartphones
may require that you download a mobile app. For those who don't want to
scan the QR codes, short URLs are provided for each individual audio and
video clip (e.g., ablle.sinauer.com/a2.1).

ABOUT THE AUDIO AND VIDEO LINKS. Most of the audio and video links
in the book are provided by researchers whose examples appear in the text, or
by the Macaulay Library at the Cornell Lab of Ornithology, drawing from their
extensive collection of nearly 7,000,000 pictures, videos, and animal sounds.
Author Biographies
Dustin Rubenstein, an Associate Professor of Ecology, Evolution, and Envi­
ronmental Biology at Columbia University, has studied animal behavior in
birds, reptiles, mammals, insects, and crustaceans for nearly 20 years through­
out Africa, Asia, Central and South America, and the Caribbean. As a leading
expert in animal social behavior and evolution, his research has been pub­
lished in top journals like Science, Nature, and PNAS, and he is co-editor of
the book Comparative Social Evolution, published in 2017. In recognition of his
research accomplishments, he has received young investigator awards from
the Animal Behavior Society, the American Ornithologists7 Union, the Society
for Behavioral Neuroendocrinology, and the University of Michigan, and been
made a Fellow of the American Ornithological Society. He was also recognized
by the National Academy of Sciences as both a Kavli Fellow for his research
accomplishments and an Education Fellow in the Sciences for his innovations
in STEM teaching.
Throughout his education and training at Dartmouth College, Cornell Uni­
versity, and the University of California, Berkeley, Rubenstein has used Animal
Behavior in his courses since its fifth edition. He seems to have learned a thing
or two, having met his wife in his undergraduate animal behavior class. From
his own work using stable isotopes to study avian migration, to studying
stress hormones and breeding behavior in birds and lizards, to examining the
genetic and epigenetic bases of reproductive behavior in insects and birds,
Rubenstein approaches the study of animal behavior in an integrative and
interdisciplinary manner. Over the past few years, he has helped lead a series
of workshops and symposia about the integrative study of behavior, a topic of
great emphasis in this new edition. His goal in taking over this classic textbook
was to highlight the modern approach to animal behavior, while maintaining
the book's timeless writing style and structure. Rubenstein thanks his wife
Kate and two children, Renna and Ian, for their patience and support over the
past few years of book writing. Many mentors helped teach and train Dustin
in animal behavior over the years, but none were more important than his
father, Daniel Rubenstein, a fellow behavioral biologist who has inspired him
since his earliest days, together observing spiders in the backyard or zebras
in Africa.
John Alcock retired in 2009 after having had an academic life full of rewards,
such as trips to Australia on sabbaticals and during leaves of absence. His
research focused on insect reproductive behavior and the many puzzles it
provides for behavioral biologists, both in his home state of Arizona as well
as in Costa Rica and Australia. Although retired, he continues to study insects,
especially in Virginia, where he lives on the family farm in the spring and
summer months. Alcock has written a number of natural history books for
a general audience, such as Sonoran Desert Spring and his most recent effort
along these lines, After the Wildfire. This book describes the account of the
environmental recovery that occurred in the ten years that followed an intense
wildfire in the mountains about an hour to the north of his home in Tempe,
 XVIII AUTHOR BIOGRAPHIES

Arizona. Now 75, Alcock enjoys being a grandfather, thanks to a son Nick and
his wife Sara who have produced two grandchildren for him (and others). His
other son Joe and his wife Satkirin are both doctors, and Joe is deeply inter­
ested in evolutionary medicine. This version of the textbook that Alcock first
wrote long, long ago has been revised by Dustin Rubenstein, a good choice to
carry the book forward as the readers of this text will shortly learn.

k I
I

Acknowledgements
For over four decades, this book has greatly benefitted from the generosity
of many colleagues who have contributed reviews, photographs, videos, and
other means of support. We are especially grateful to the reviewers for this
eleventh edition, whose suggestions improved the structure, content, and tone
of many of the chapters. The reviewers for this edition include:

Patrick Abbot Vanderbilt University

Andres Bendesky Columbia University
Kirsten M. Bohn Johns Hopkins University
Jennifer S. Borgo Coker College
Patty L.R. Brennan Mount Holyoke College
Molly Cummings The University of Texas at Austin
Ben Dantzer University of Michigan
Harold Gouzoules Emory University
Sarah Guindre-Parker University of Guelph
Kristina M. Hannam State University of NY-Geneseo
Margret I. Hatch Penn State Worthington Scranton
Mark E. Hauber City University of New York
Hans Hofmann University of Texas at Austin
Victoria Ingalls Marist College
Roland Kays North Carolina Museum of Natural Sciences
Darcy Kelley Columbia University
Walt Koenig Cornell University
Rafael Maia Columbia University
Heather Masonjones University of Tampa
Ronald L. Mumme Allegheny College
Wiline M. Pangle Central Michigan University
Kimberly N. Russell Rutgers University
D. Kim Sawrey University of North Carolina Wilmington
Joseph Sisneros University of Washington
Nancy G. Solomon Miami University
Elizabeth Tibbetts University of Michigan
Sarah Woolley Columbia University
Tim Wright New Mexico State University
Ken Yasukawa Beloit College

We are also indebted to our wonderful colleagues at Sinauer Associates. With­

out their dedication, patience, and careful oversight (and occasional prodding),
this book—as well as the ten previous editions—would never have made it to
print. Hie team that worked on this edition included:

Andrew D. Sinauer, Publisher Joanne Delphia, Book Designer

Rachel Meyers, Acquisitions Editor Mark Siddall, Photo Researcher
Martha Lorantos, Production Editor Elizabeth Pierson, Copy Editor
Chris Small, Production Manager
Beth Roberge Friedrichs, Production
Specialist
Media and
Supplements
to accompany Animal Behavior,
Eleventh Edition

eBook
(ISBN 9-781-60535-770-6)
Animal Behavior, Eleventh Edition, is available as an eBook, in several different
formats, including RedShelf, VitalSource, and Chegg. All major mobile devices
are supported.

For the Student

Companion Websito (oup.com/us/rubensteinlle)
The Animal Behavior, Eleventh Edition, Companion Website includes a vari­
ety of study and review aids—all available at no cost to the student. The site
includes the following:
• Chapter Summaries give the student a thorough review of each
chapter's content.
• Audio and Video Links provide a set of online sites and resources
relevant to each chapter.
• The Glossary provides definitions for all textbook bolded terms.

Student Lab Manual

This electronic lab manual walks students through the scientific process to
improve their fluency with hypothesis development, observing and quantify­
ing animal behavior, statistical analysis, and data presentation.

For the Instructor

Ancillary Resource Center (oup-arc.com)
The Ancillary Resource Center provides instructors using Animal Behavior with
a wealth of resources for use in course planning, lecture development, and
assessment. Contents include:
TEXTBOOK FIGURES AND TABLES
All the figures and tables from the textbook are provided in JPEG format,
reformatted for optimal readability, with complex figures provided in both
whole and split formats.

■
 MEDIA AND SUPPLEMENTS XXI

POWERPOINT RESOURCES
This presentation includes all of the figures and tables from the chapter, with
titles on each slide and complete captions in the Notes field.
INSTRUCTOR’S MANUAL AND TEST BANK
These resources facilitate the preparation of lectures, quizzes, and exams.
Contents include:
° Answers to the discussion questions in the textbook
® A list of films and documentaries about animal behavior, for use in
the classroom
• A test bank, consisting of a range of multiple-choice and short
answer questions drawn from textbook content, along with writing
assignments to engage students in deeper thinking.

Learning the Skills of Research:

Animal Behavior Exercises in the Laboratory and Field
Edited by Elizabeth M. Jakob and Margaret Hodge
Students learn best about the process of science by carrying out projects from
start to finish. Animal behavior laboratory classes are particularly well suited
for independent student research, as high-quality projects can be conducted
with simple materials and in a variety of environments. The exercises in this
electronic lab manual are geared to helping students learn about all stages of
the scientific process: hypothesis development, observing and quantifying ani­
mal behavior, statistical analysis, and data presentation. Additional exercises
allow the students to practice these skills, with topics ranging from habitat
selection in isopods to human navigation. The manuals provide both student
and instructor documentation. Data sheets and other supplementary material
are offered in editable formats that instructors can modify as desired.

To learn more about any of these resources, or to get access, please contact your
OUP representative.
Natural Selection and the
Evolution of Behavior 5
The Cost-Benefit Approach to
Behavioral Biology
The Levels of Analysis
The Integrative Study of
Animal Behavior
Approaches to Studying
Behavior 13
The Adaptive Basis of Behavior:
Mobbing in Gulls
The Science of Animal Behavior
n Introduction to
Animal Behavior
f you’ve ever been stung by a honey bee (Apis mellifera), you
know that animals can behave in ways that seem illogical to hu­
mans. After all, just a few minutes after stinging you, that female
honey bee will die. Why would an individual kill itself to temporar­
ily hurt you? Although we will not fully answer this question until
Chapter 12, the solution lies deep in the portion of the stinging
bee’s genetic code that it shares with its hive mates. As you will
see in this chapter, behavioral theory gives us a framework within
which to understand this and other seemingly paradoxical be­
haviors. As a student of animal behavior, you will use this book
to begin exploring the natural world and examine how and why
the diversity of animals that live in it behave in the ways they do.
 1

4 CHAPTER 1

Chapter 1 opening photograph The discipline of animal behavior is vibrant and growing rapidly, thanks to thou­
- by duncan1890/E+/Getty sands of behavioral biologists who are exploring everything from the genetics of bird
Images.
song to why women find men with robust chins attractive. A major reason why the
field is so active and broad ranging has to do with a book published more than 150
years ago, Charles Darwin's On the Origin of Species (Darwin 1859). In this book, Dar­
win (FIGURE 1.1) introduced the concept of natural selection, which argues that
living species are the product of an unguided, unconscious process of reproductive
competition among their ancestors. Natural selection theory provides the guiding
principle for studying animal behavior, as well as most of biology more generally.
I Knowing that animal behavior, like every other aspect of living things, has a history
guided by natural selection is hugely important. An understanding of evolutionary
theory gives us a scientific starting point when we set out to determine why animals
do the things they do—and why they have the genetic, developmental, sensory, neu­
ronal, physiological, and hormonal mechanisms that make these behavioral abilities
possible. To truly understand why a bee will kill itself to sting you, you must approach
the study of animal behavior from an evolutionary perspective, from the perspective
of Darwin himself. As the evolutionary biologist Theodosius Dobzhansky once said,
"Nothing in biology makes sense except in the light of evolution" (Dobzhansky 1973).
We hope to convince you that Dobzhansky was right when it comes to animal behav­
ior as well. If we succeed, you will come to understand the appeal of the evolutionary
approach to animal behavior, as it helps scientists identify interesting subjects worthy
of explanation, steers them toward hypotheses suitable for testing, and ultimately
produces robust conclusions about the validity of certain hypotheses.
Darwin's influence continues strongly to this day, which is why this book was titled
Animal Behavior: An Evolutionary Approach for its first ten editions. Over the editions,
we have explored the assortment of behaviors that animals exhibit and
discuss not only how and why they evolved, but why they evolved
in one species and not another, or in one sex and not the other. In this
eleventh edition, we extend the evolutionary approach emphasized
by the subtitle in the book's prior editions and take a more integra­
tive approach. The concept of evolution by natural selection remains
our guide, but we will also weave into our discussion topics such as
epigenetics, development, and ecology, as well as tools such as game
theory and molecular genetics. Indeed, as you will see in the following
chapters, the study of animal behavior has itself evolved since Darwin
made his observations about natural selection more than 150 years ago.
In fact, the study of animal behavior has changed considerably since
the first edition of this book was published more than 40 years ago.
Therefore, in this edition, we go beyond an evolutionary approach to
the study of animal behavior. We also consider an ecological approach.
A mechanistic approach. A developmental approach. In short, we take
an integrative approach to the study of animal behavior (Rubenstein
and Hofmann 2015, Hofmann et al. 2016), discussing how the inter­
action among genes, development, and the environment shapes the
evolution of animal behaviors. This means that students of animal
behavior today may need to use the material learned in a biochemistry
or molecular biology class just as much as they do the material learned
in an ecology class. We illustrate the ways, for example, that genomics
and neurobiology are changing how we study animal behavior in a
FIGURE 1.1 Charles Darwin. Taken in
manner that Darwin could not have imagined. And we demonstrate
1831 by Herbert Rose Barraud (1845-1896),
this is thought to be the last photograph of how new phylogenetic comparative methods are enabling us to use
Darwin before his death. The original is in The data collected from dozens to thousands of species to test existing, as
Huntington Library in San Marino, California. well as generate novel, hypotheses.

a
 AN INTRODUCTION TO ANIMAL BEHAVIOR

But first we introduce you to the fascinating field of animal behavior in this chap­
ter. We begin with a detailed discussion of how natural selection governs the evolu­
tion of behavior. We introduce adaptationist thinking and the gene's eye view of
behavioral trait evolution. We discuss the cost-benefit approach to behavioral biol­
ogy, as well as the critical role that hypothesis testing plays in the scientific method.
Ultimately, we argue that an integrative approach to studying animal behavior is
essential for understanding how and why behaviors evolve, a topic we will explore
in greater detail in Chapter 2 and then throughout the rest of the book.

Natural Selection and

the Evolution of Behavior
When biologists ask questions about the behavior of animals, they are guided by
Darwin's theory of evolution by natural selection. Darwin was puzzled by the fact
that living organisms could increase their numbers geometrically, but that most
didn't. Even in bacteria, which can reproduce rapidly and efficiently, some individu­
als replicate more than others. So which individuals reproduce more, and why? As
Darwin came to realize after a lifetime of observing animals in their natural habitat,
if in the past some individuals left more descendants than others, then these repro-
ductively successful individuals would inevitably gradually reshape their species
in their image. The logic of natural selection is such that evolutionary change is
inevitable if just three conditions are met:
1. Variation, with members of a population differing in a particular
characteristic
2. Differential reproductive success, with some individuals with particular
characteristics having more offspring than others
3. Heredity, with parents able to pass on those characteristics to their offspring
If there is variation within a species (and there almost always is), if some of that
variation is heritable and passed from parents to offspring, and if some of those
individuals consistently reproduce more successfully than others, then the increased
abundance of living descendants of the more successful types will gradually change
the species. Over time, the "old" population—it is important to remember that natu­
ral selection acts on populations of individuals—evolves into one whose members
possess the characteristics (or traits) that were associated with successful reproduc­
tion in the past. How successful an individual is at passing on its heritable traits to
the next generation is referred to as fitness. As you will see throughout this book,
fitness—which depends on both survival and reproduction—forms the foundation
for understanding which traits are likely to evolve via natural selection and become
more or less common in a population. After all, to reproduce and pass one's traits
on to one's offspring, an individual has to survive long enough to breed.
Darwin not only laid out the logic of his theory clearly, he also provided abun­
dant evidence that heritable variation in traits is common within species, and that
high rates of mortality are also the rule. Indeed, the conditions necessary and suf­
ficient for evolutionary change by natural selection to occur are present in all living
things, a point that Darwin demonstrated by showing that people could cause dogs
and pigeons to evolve by selectively breeding those individuals with traits that the
breeders wanted in future generations of their domesticated animals. And although
it was a heretical concept in Darwin's Victorian England, we now know that people,
like all other organisms, also evolve by natural selection. For example, the ability
of many humans to continue to digest lactose, the dominant sugar in milk, after

__
6 CHAPTER 1

childhood appeared around the time our ancestors domesticated livestock and has
been hypothesized to have been strongly favored by natural selection because it
allowed some individuals to survive on a milk diet.
We call the traits associated with successful survival and reproduction, and on
which natural selection acts, adaptations. Figuring out exactly how a putative
adaptation contributes to the reproductive success of individuals is perhaps the
central goal for most behavioral biologists, some of whom are happy to be known as
"adaptationists." Certainly Darwin would have agreed with the way adaptationists
think about evolution, especially with respect to the hereditary foundation of evo­
lutionary theory. Although Darwin himself knew nothing about genes, regions of
DNA that encode traits, we now can reconfigure his argument to deal with selection
at the level of the gene. Just as adaptations that increase the reproductive success of
individuals will spread through populations over time, so too will the hereditary
basis for these attributes. Because genes can be present in populations in different
forms known as alleles, those alleles that contribute to traits linked to individual
reproductive success will become more common over time, whereas those associ­
ated with reproductive failure will eventually disappear from the population and
perhaps even the genome. In evolutionary biology, we refer to traits as pheno­
types (aspects of an individual that arise from an interaction of the individual's
genes with its environment), and to the set of alleles underlying the development of
those traits as genotypes. The genetic basis of most complex phenotypes—includ­
ing most behaviors—is rarely known in full, and when it is, it is typically only in
model organisms. (Although as you will see in subsequent chapters, this is changing
and scientists are now capable of identifying the hereditary basis of a trait in almost
any organism.) Linking phenotypes to genotypes is not only an essential part of
modem molecular and evolutionary biology, but with the powerful new tools and
approaches from these disciplines being applied to non-model organisms in their
natural habitats, it is also becoming crucial to behavioral biology as well.
It is critical to recognize three points about this so-called gene-centered, or gene's
eye, view of evolution by natural selection:
1. Only genes replicate themselves; organisms do not. Instead, organisms—or
groups of organisms—are vehicles within which replicators (genes) travel, a
point we will discuss further in Chapter 12 (Dawkins 1989). Adaptive evolu­
tion therefore occurs through the differential survival of competing genes,
which increases the frequency of those alleles whose phenotypic effects
promote their own propagation.
2. Evolution is not natural selection. Evolution is gene frequency change within
a population. Natural selection is one of several causes of evolution, as are
mutation, migration, and genetic drift.
3. Natural selection is not guided by anything or anyone. Selection is not
"trying" to do anything. Instead, it is the individuals that reproduce more
that cause a population or species to evolve over time, a process that Darwin
called descent with modification.
Notice also that the only kinds of heritable traits that will become more common
in a species are those that promote individual reproductive success, which do not
necessarily benefit the species as a whole. Although "for the good of the species"
arguments were often made by biologists not so long ago, it is entirely possible for
adaptations (and particular alleles) to spread through populations even if they do
nothing to perpetuate the species (see Chapter 12). Indeed, traits and alleles that are
harmful to group survival in the long run can still be favored by natural selection.
Yet as we will discuss later in this book, there are also special cases when selection
 AN INTRODUCTION TO ANIMAL BEHAVIOR 7

acts on traits that do benefit others, even seemingly at the expense of the individu­
als that possess those traits.

The Cost-Benefit Approach to Behavioral Biology

To help establish why a particular behavior has evolved, behavioral biologists must
propose a hypothesis, or an explanation based on limited evidence. Of course, for
most behaviors there may be multiple potential explanations, which requires pos­
ing multiple hypotheses. When only one of a series of competing hypotheses could
explain a given behavior, we call these alternative hypotheses. In contrast, when
multiple hypotheses could apply to a given behavior, we refer to them as non-
mutually exclusive hypotheses. Throughout this book, we highlight hypotheses
in blue in the text, and also define them in the page margins. When there are multiple
hypotheses to explain a behavior, we group them together in a margin box and note
whether they are alternative or non-mutually exclusive hypotheses.
Once a hypothesis has been generated, researchers can use it to make a prediction,
or expectation that should follow if the hypothesis is true. Predictions allow for the
discrimination among competing hypotheses. Generating, falsifying, and then gen­
erating new hypotheses and predictions is a natural part of the scientific method.
When behavioral biologists set out to test hypotheses and predictions related
to a potentially adaptive behavior, they often consider the costs and benefits of
that behavior; they take a cost-benefit approach to the study of animal behavior.
When they speak of costs, behavioral biologists are talking about fitness costs,
the negative effects of a trait on the number of surviving offspring produced by an
individual or a reduction in the number of copies of its alleles that it contributes
to the next generation. When they speak of fitness benefits, they are referring
to the positive effects of a trait on reproductive (and genetic) success. Fitness costs
and benefits are the units that behavioral biologists use to study adaptations and
the process of evolution by natural selection. Most behavioral biologists study
traits that they assume are adaptations, an assumption that they make in order to
test specific hypotheses about the possible adaptive value, or the contribution
to fitness, of the characteristic of interest. Recall that an adaptation is a heritable
trait that has spread or is spreading by natural selection, and has replaced or is
replacing any alternative traits in the population or species. Such an attribute has
a better ratio of fitness benefits to costs than the alternative forms of this trait that
have occurred in the species. Thus, studying why a behavioral adaptation has
evolved depends on measuring the potential reproductive costs and benefits to
the individual adopting that behavior.
Since fitness is an abstract term, behavioral biologists often have to settle for an
indicator or correlate of reproductive success when they attempt to measure fitness.
In the chapters that follow, the terms fitness, reproductive success, and genetic success
are often used more or less interchangeably when referring to such indicators as
offspring survival, the number of young that survive to fledging, the number of
mates inseminated, or even more indirectly, the quantity of food ingested per unit
of time, the ability to acquire a breeding territory, and so on. These proxies give us
a currency with which to measure fitness costs and benefits and ultimately to test
hypotheses underlying the evolution of a range of animal behaviors.
Using natural selection theory and the cost-benefit approach helps us identify
why behaviors evolve, including seeming anomalies that require explanation such
as why traits that appear to reduce rather than raise an individual's reproductive
success persist or even spread through a population. As we discussed at the outset,
a honey bee sting is a good example of a behavioral trait that at first seems counter­
intuitive, or even maladaptive for the individual doing the stinging. (In Chapter 12
8 CHAPTER 1

FIGURE 1.2 A male Hon

(Panthera led) carrying a
young lion cub that he has
killed. Because infanticide
fh
&
, ..M
by males occurs in several
different species under similar tt*' \ : - ^
social circumstances, the be­
havior is likely to be adaptive. TS^i. ■
(Photograph © Laura Romin
and Larry Dalton/Alamy Stock
Photo.)

we will explore why animals perform self-sacrificial acts such as this.) Throughout
the book, we will refer to these challenges to evolutionary theory as Darwinian
puzzles. Biologists deal with these puzzles by developing possible hypotheses
based on natural selection theory for how the trait might actually help individuals
reproduce and pass on their genes. As an illustration of this approach, we describe
in BOX 1.1 an example of a behavior that has long puzzled behavioral biologists:
infanticide (FIGURE 1.2).

The Levels of Analysis

Now that we have discussed how traits—including behavioral ones—evolve via
natural selection, let's consider what a behavioral trait is and how we study it.
Although "behavior" seems like it should be an easy term to define, it actually
means different things to different people, particularly those in different fields. You
would think that behavioral biologists could at least agree on what a behavior is,
but they don't. Daniel Levitis and colleagues posed a seemingly simple question—
"What is animal behavior?"—to nearly 175 behavioral biologists at a series of sci­
entific society meetings, and much to their surprise, there was no consensus on
how behavioral biologists define animal behavior. Levitis and colleagues suggested
defining behavior as the internally coordinated responses (actions or inactions) of
whole living organisms (individuals or groups) to internal and/or external stimuli,
excluding responses more easily understood as developmental changes (Levitis et
al. 2009). This definition focuses largely on intrinsic factors. What if we gave you a
different definition, something like this: behavior describes the way an individual
acts or interacts with others or the environment? This definition has nothing to do
with the internal processes of an individual. Instead, this definition emphasizes
interactions between individuals or between an individual and its environment.
Why do these definitions of behavior differ so much?
Part of the reason why behavioral biologists define behavior so differently is
that animal behavior is a wide and varied discipline that has itself evolved over
time. The field of animal behavior has inherited traditions from ethology and the

I
 AN INTRODUCTION TO ANIMAL BEHAVIOR 9

BOX
1.1 DARWINIAN PUZZLE

I Natural selection and infanticide in primates

Hanuman langurs (Semnopithecus entellus) aggressive behavior, and infanticide was

are monkeys that live in groups of several simply a maladaptive result (Curtin and
females and their offspring, accompanied Dolhinow 1978).
by one or a few adult males (FIGURE A). In (Continued)
the course of a long-term research project
in India, male Hanuman langurs were
(A)
seen attacking and sometimes even
killing the very young infants of females
in their own group. The puzzle here is
obvious: how can it be adaptive for a male
langur to harm the offspring of females
in his group, particularly since attacking
males are sometimes injured by mothers
defending their babies (FIGURE B)?
Some primatologists have argued that the
infanticidal behavior of these males was
not adaptive but was instead the aberrant
aggressive response by males to the
overpopulation and crowding that occurred
when langurs came together to be fed
by Indian villagers. According to these FIGURE A Hanuman langur females and
observers, overcrowding caused abnormal offspring. Males fight to monopolize sexual access
to the females in groups such as this one. (Photo­
graph © Heini Wehrle/AGE Fotostock.)

(B)

FIGURE B Male langurs commit infanticide. (Left)

A nursing baby langur that has been paralyzed by a male
langur’s bite to the spine (note the open wound). This infant
was attacked repeatedly over a period of weeks, losing an
eye and finally its life at age 18 months. (Right) An infant­
killing male langur flees from an aggressive protective female
belonging to the band he is attempting to join. (Left, photo­
graph by Carola Borries; right, photograph by Volker Som­
mer, from Sommer 1987.)
 lO CHAPTER 1

BOX DARWINIAN PUZZLE (continued)

1.1

Behavioral biologist Sarah Hrdy used natural
selection theory to try to solve the puzzle of
infanticide in a different way, namely by asking
whether the killer males were behaving in a
reproductively advantageous manner (Hrdy
1977). She reasoned that by committing
infanticide, the males might cause the baby­
less females to resume ovulating, which
otherwise does not happen for several years
in females that retain and nurse their infants.
Once females began ovulating, the males could
then mate with them. Hrdy tried to explain
how infanticide might have spread through
Hanuman langur populations in the past as a
reproduction-enhancing tactic for individual
males. This infanticide hypothesis leads
to several expectations, of which the most
important is the prediction that males will not kill
their own progeny, but will instead focus their
attacks on the offspring of other males. This
prediction in turn generates the expectation
that infanticide will be linked to the arrival of
a new male or males into a group of females,
with the associated ejection of the father or
fathers of any baby langurs in the group. In
cases of male turnover in groups, the new
males could father offspring more quickly if
they first killed the existing infants. Females
that lose their infants do resume ovulating,
which enables the new males in the group
to father replacement offspring. Since these
predictions have been shown to be correct
for this species (Borries et al. 1999) as well
as for some other primates (Beehner and
Bergman 2008, Lyon et al. 2011) and for various
carnivores, horses, rodents, and even a bat
(Kndrnschild et al. 2011), we can conclude that
infanticide in Hanuman langurs is indeed an
adaptation, the product of natural selection.
infanticide hypothesis Infanticide is a
reproduction-enhancing tactic practiced by
males.

V Thinking Outside the Box.....................

Given what you just read about the gene’s
eye view of evolution by natural selection, can
you rephrase the infanticide hypothesis using
this terminology? In other words, consider
infanticide from a gene’s perspective and
discuss why it could be a beneficial behavior
from the perspective of a gene that underlies
that behavior. Would a gene for infanticidal
behavior be more or less likely to be passed
on than a gene that does not promote the
killing of young?

observation of animals in the wild, as well as those from experimental psychology

where experiments were generally conducted in the lab. As we said at the outset,
animal behavior is becoming increasingly integrative and continues to evolve as
a discipline to this day. Some scientists tend to take a more mechanistic or devel­
opmental approach, as our first definition implied. Still other scientists take an
adaptive or evolutionary approach, as our second definition suggested. Indeed,
these ultimate causes of behavior provide us with a perspective on the adaptive
value and the evolutionary history of a trait of interest. However, there is much
more to behavioral research than studies of adaptation and evolutionary history.
As the behavioral biologist Nikolaas Tinbergen pointed out long ago, biologists
also need to study how a behavior develops over the lifetime of an individual and
how an animal's physiological systems make behaving possible if they are to really
understand the evolution of a behavior (Tinbergen 1963). Because the mechanisms
that underlie development and neurophysiology act within the life span of indi­
viduals, they are considered to be immediate and mechanistic, or what we call
I

L
Another random document with
no related content on Scribd:
 grace, her form all of beauty to me who opposite sat and was
watching her dextrous fingers.
The manufacture of flax into linen material was ever felt to be of
vast importance, and was encouraged by legislation from earliest
colonial days, but it received a fresh impulse in New England
through the immigration of about one hundred Irish families from
Londonderry. They settled in New Hampshire on the Merrimac about
1719. They spun and wove by hand, but with far more skill than
prevailed among those English settlers who had already become
Americans. They established a manufactory according to Irish
methods, and attempts at a similar establishment were made in
Boston. There was much public excitement over spinning. Women,
rich as well as poor, appeared on Boston Common with their wheels,
thus making spinning a popular holiday recreation. A brick building
was erected as a spinning-school, and a tax was placed in 1737 to
support it. But this was not an industrial success, the excitement died
out, the public spinning-school lost its ephemeral popularity, and the
wheel became again simply a domestic duty and pride.
For many years after this, housewives had everywhere flax and
hemp to spin and weave in their homes, and the preparation of these
staples seems to us to-day a monumental labor. On almost every
farm might be seen a patch of the pretty flax, ripening for the hard
work of pulling, rippling, rotting, breaking, swingling, and combing,
which all had to be done before it came to the women’s hands for
spinning. The seed was sown broad-cast, and allowed to grow till the
bobs or bolls were ripe. The flax was then pulled and spread neatly
in rows to dry. This work could be done by boys. Then men whipped
or threshed or rippled out all the seed to use for meal; afterwards the
flax stalks were allowed to lie for some time in water until the shives
were thoroughly rotten, when they were cleaned and once more
thoroughly dried and tied in bundles. Then came work for strong
men, to break the flax on the ponderous flaxbreak, to get out the
hard “hexe” or “bun,” and to swingle it with a swingle knife, which
was somewhat like a wooden dagger. Active men could swingle forty
pounds a day on the swingling-board. It was then hetchelled or
combed or hackled by the housewife, and thus the rough tow was
gotten out, when it was straightened and made ready for the spruce
distaff, round which it was finally wrapped. The hatchelling was
tedious work and irritating to the lungs, for the air was filled with the
fluffy particles which penetrated everywhere. The thread was then
spun on a “little wheel.” It was thought that to spin two double skeins
of linen, or four double skeins of tow, or to weave six yards of linen,
was a good day’s work. For a week’s work a girl received fifty cents
and “her keep.” She thus got less than a cent and a half a yard for
weaving. The skeins of linen thread went through many tedious
processes of washing and bleaching before being ready for weaving;
and after the cloth was woven it was “bucked” in a strong lye, time
and time again, and washed out an equal number of times. Then it
was “belted” with a maple beetle on a smooth, flat stone; then
washed and spread out to bleach in the pure sunlight. Sometimes
the thread, after being spun and woven, had been washed and
belted a score of times ere it was deemed white and soft enough to
use. The little girls could spin the “swingling tow” into coarse twine,
and the older ones make “all tow” and “tow and linen” and “harden”
stuffs to sell.
To show the various duties attending the manufacture of these
domestic textiles by a Boston woman of intelligence and social
standing, as late as 1788, let me quote a few entries from the diary
of the wife of Col. John May:—
A large kettle of yarn to attend upon. Lucretia and self rinse
our through many waters, get out, dry, attend to, bring in, do
up and sort 110 score of yarn, this with baking and ironing.
Went to hackling flax.
Rose early to help Ruth warp and put a piece in the loom.
Baking and hackling yarn. A long web of tow to whiten and
weave.
The wringing out of this linen yarn was most exhausting, and the
rinsing in various waters was no simple matter in those days, for the
water did not conveniently run into the houses through pipes and
conduits, but had to be laboriously carried in pailfuls from a pump, or
more frequently raised in a bucket from a well.
I am always touched, when handling the homespun linens of olden
times, with a sense that the vitality and strength of those enduring
women, through the many tedious and exhausting processes which
they had bestowed, were woven into the warp and woof with the flax,
and gave to the old webs of linen their permanence and their
beautiful texture. How firm they are, and how lustrous! And how
exquisitely quaint and fine are their designs; sometimes even
Scriptural designs and lessons are woven into them. They are,
indeed, a beautiful expression of old-time home and farm life. With
their close-woven, honest threads runs this finer beauty, which may
be impalpable and imperceptible to a stranger, but which to me is
real and ever-present, and puts me truly in touch with the life of my
forbears. But, alas, it is through intuition we must learn of this old-
time home life, for it has vanished from our sight, and much that is
beautiful and good has vanished with it.
The associations of the kitchen fireside that linger in the hearts of
those who are now old can find no counterpart in our domestic
surroundings to-day. The welcome cheer of the open fire, which
graced and beautified even the humblest room, is lost forever with
the close gatherings of the family, the household occupations, the
homespun industries which formed and imprinted in the mind of
every child the picture of a home.
 Transcriber’s Notes
Minor punctuation errors have been silently corrected.
Page 100: “take the the case” changed to “take the case”
Page 162: “promply sailed” changed to “promptly sailed”
Page 302: “was was set outside” changed to “was set outside”
Spelling and punctuation quoted from original sources has been left as-is.
*** END OF THE PROJECT GUTENBERG EBOOK COLONIAL
DAMES AND GOOD WIVES ***

Updated editions will replace the previous one—the old editions

will be renamed.

Creating the works from print editions not protected by U.S.

copyright law means that no one owns a United States copyright
in these works, so the Foundation (and you!) can copy and
distribute it in the United States without permission and without
paying copyright royalties. Special rules, set forth in the General
Terms of Use part of this license, apply to copying and
distributing Project Gutenberg™ electronic works to protect the
PROJECT GUTENBERG™ concept and trademark. Project
Gutenberg is a registered trademark, and may not be used if
you charge for an eBook, except by following the terms of the
trademark license, including paying royalties for use of the
Project Gutenberg trademark. If you do not charge anything for
copies of this eBook, complying with the trademark license is
very easy. You may use this eBook for nearly any purpose such
as creation of derivative works, reports, performances and
research. Project Gutenberg eBooks may be modified and
printed and given away—you may do practically ANYTHING in
the United States with eBooks not protected by U.S. copyright
law. Redistribution is subject to the trademark license, especially
commercial redistribution.

START: FULL LICENSE

THE FULL PROJECT GUTENBERG LICENSE
 PLEASE READ THIS BEFORE YOU DISTRIBUTE OR USE THIS WORK

To protect the Project Gutenberg™ mission of promoting the

free distribution of electronic works, by using or distributing this
work (or any other work associated in any way with the phrase
“Project Gutenberg”), you agree to comply with all the terms of
the Full Project Gutenberg™ License available with this file or
online at www.gutenberg.org/license.

Section 1. General Terms of Use and

Redistributing Project Gutenberg™
electronic works
1.A. By reading or using any part of this Project Gutenberg™
electronic work, you indicate that you have read, understand,
agree to and accept all the terms of this license and intellectual
property (trademark/copyright) agreement. If you do not agree to
abide by all the terms of this agreement, you must cease using
and return or destroy all copies of Project Gutenberg™
electronic works in your possession. If you paid a fee for
obtaining a copy of or access to a Project Gutenberg™
electronic work and you do not agree to be bound by the terms
of this agreement, you may obtain a refund from the person or
entity to whom you paid the fee as set forth in paragraph 1.E.8.

1.B. “Project Gutenberg” is a registered trademark. It may only

be used on or associated in any way with an electronic work by
people who agree to be bound by the terms of this agreement.
There are a few things that you can do with most Project
Gutenberg™ electronic works even without complying with the
full terms of this agreement. See paragraph 1.C below. There
are a lot of things you can do with Project Gutenberg™
electronic works if you follow the terms of this agreement and
help preserve free future access to Project Gutenberg™
electronic works. See paragraph 1.E below.
1.C. The Project Gutenberg Literary Archive Foundation (“the
Foundation” or PGLAF), owns a compilation copyright in the
collection of Project Gutenberg™ electronic works. Nearly all the
individual works in the collection are in the public domain in the
United States. If an individual work is unprotected by copyright
law in the United States and you are located in the United
States, we do not claim a right to prevent you from copying,
distributing, performing, displaying or creating derivative works
based on the work as long as all references to Project
Gutenberg are removed. Of course, we hope that you will
support the Project Gutenberg™ mission of promoting free
access to electronic works by freely sharing Project
Gutenberg™ works in compliance with the terms of this
agreement for keeping the Project Gutenberg™ name
associated with the work. You can easily comply with the terms
of this agreement by keeping this work in the same format with
its attached full Project Gutenberg™ License when you share it
without charge with others.

1.D. The copyright laws of the place where you are located also
govern what you can do with this work. Copyright laws in most
countries are in a constant state of change. If you are outside
the United States, check the laws of your country in addition to
the terms of this agreement before downloading, copying,
displaying, performing, distributing or creating derivative works
based on this work or any other Project Gutenberg™ work. The
Foundation makes no representations concerning the copyright
status of any work in any country other than the United States.

1.E. Unless you have removed all references to Project

Gutenberg:

1.E.1. The following sentence, with active links to, or other

immediate access to, the full Project Gutenberg™ License must
appear prominently whenever any copy of a Project
Gutenberg™ work (any work on which the phrase “Project
Gutenberg” appears, or with which the phrase “Project
Gutenberg” is associated) is accessed, displayed, performed,
viewed, copied or distributed:

This eBook is for the use of anyone anywhere in the United

States and most other parts of the world at no cost and with
almost no restrictions whatsoever. You may copy it, give it
away or re-use it under the terms of the Project Gutenberg
License included with this eBook or online at
www.gutenberg.org. If you are not located in the United
States, you will have to check the laws of the country where
you are located before using this eBook.

1.E.2. If an individual Project Gutenberg™ electronic work is

derived from texts not protected by U.S. copyright law (does not
contain a notice indicating that it is posted with permission of the
copyright holder), the work can be copied and distributed to
anyone in the United States without paying any fees or charges.
If you are redistributing or providing access to a work with the
phrase “Project Gutenberg” associated with or appearing on the
work, you must comply either with the requirements of
paragraphs 1.E.1 through 1.E.7 or obtain permission for the use
of the work and the Project Gutenberg™ trademark as set forth
in paragraphs 1.E.8 or 1.E.9.

1.E.3. If an individual Project Gutenberg™ electronic work is

posted with the permission of the copyright holder, your use and
distribution must comply with both paragraphs 1.E.1 through
1.E.7 and any additional terms imposed by the copyright holder.
Additional terms will be linked to the Project Gutenberg™
License for all works posted with the permission of the copyright
holder found at the beginning of this work.

1.E.4. Do not unlink or detach or remove the full Project

Gutenberg™ License terms from this work, or any files
containing a part of this work or any other work associated with
Project Gutenberg™.
 1.E.5. Do not copy, display, perform, distribute or redistribute
this electronic work, or any part of this electronic work, without
prominently displaying the sentence set forth in paragraph 1.E.1
with active links or immediate access to the full terms of the
Project Gutenberg™ License.

1.E.6. You may convert to and distribute this work in any binary,
compressed, marked up, nonproprietary or proprietary form,
including any word processing or hypertext form. However, if
you provide access to or distribute copies of a Project
Gutenberg™ work in a format other than “Plain Vanilla ASCII” or
other format used in the official version posted on the official
Project Gutenberg™ website (www.gutenberg.org), you must, at
no additional cost, fee or expense to the user, provide a copy, a
means of exporting a copy, or a means of obtaining a copy upon
request, of the work in its original “Plain Vanilla ASCII” or other
form. Any alternate format must include the full Project
Gutenberg™ License as specified in paragraph 1.E.1.

1.E.7. Do not charge a fee for access to, viewing, displaying,

performing, copying or distributing any Project Gutenberg™
works unless you comply with paragraph 1.E.8 or 1.E.9.

1.E.8. You may charge a reasonable fee for copies of or

providing access to or distributing Project Gutenberg™
electronic works provided that:

• You pay a royalty fee of 20% of the gross profits you derive from
the use of Project Gutenberg™ works calculated using the
method you already use to calculate your applicable taxes. The
fee is owed to the owner of the Project Gutenberg™ trademark,
but he has agreed to donate royalties under this paragraph to
the Project Gutenberg Literary Archive Foundation. Royalty
payments must be paid within 60 days following each date on
which you prepare (or are legally required to prepare) your
periodic tax returns. Royalty payments should be clearly marked
as such and sent to the Project Gutenberg Literary Archive
Foundation at the address specified in Section 4, “Information
 about donations to the Project Gutenberg Literary Archive
Foundation.”

• You provide a full refund of any money paid by a user who

notifies you in writing (or by e-mail) within 30 days of receipt that
s/he does not agree to the terms of the full Project Gutenberg™
License. You must require such a user to return or destroy all
copies of the works possessed in a physical medium and
discontinue all use of and all access to other copies of Project
Gutenberg™ works.

• You provide, in accordance with paragraph 1.F.3, a full refund of

any money paid for a work or a replacement copy, if a defect in
the electronic work is discovered and reported to you within 90
days of receipt of the work.

• You comply with all other terms of this agreement for free
distribution of Project Gutenberg™ works.

1.E.9. If you wish to charge a fee or distribute a Project

Gutenberg™ electronic work or group of works on different
terms than are set forth in this agreement, you must obtain
permission in writing from the Project Gutenberg Literary
Archive Foundation, the manager of the Project Gutenberg™
trademark. Contact the Foundation as set forth in Section 3
below.

1.F.

1.F.1. Project Gutenberg volunteers and employees expend

considerable effort to identify, do copyright research on,
transcribe and proofread works not protected by U.S. copyright
law in creating the Project Gutenberg™ collection. Despite
these efforts, Project Gutenberg™ electronic works, and the
medium on which they may be stored, may contain “Defects,”
such as, but not limited to, incomplete, inaccurate or corrupt
data, transcription errors, a copyright or other intellectual
property infringement, a defective or damaged disk or other
medium, a computer virus, or computer codes that damage or
cannot be read by your equipment.

1.F.2. LIMITED WARRANTY, DISCLAIMER OF DAMAGES -

Except for the “Right of Replacement or Refund” described in
paragraph 1.F.3, the Project Gutenberg Literary Archive
Foundation, the owner of the Project Gutenberg™ trademark,
and any other party distributing a Project Gutenberg™ electronic
work under this agreement, disclaim all liability to you for
damages, costs and expenses, including legal fees. YOU
AGREE THAT YOU HAVE NO REMEDIES FOR NEGLIGENCE,
STRICT LIABILITY, BREACH OF WARRANTY OR BREACH
OF CONTRACT EXCEPT THOSE PROVIDED IN PARAGRAPH
1.F.3. YOU AGREE THAT THE FOUNDATION, THE
TRADEMARK OWNER, AND ANY DISTRIBUTOR UNDER
THIS AGREEMENT WILL NOT BE LIABLE TO YOU FOR
ACTUAL, DIRECT, INDIRECT, CONSEQUENTIAL, PUNITIVE
OR INCIDENTAL DAMAGES EVEN IF YOU GIVE NOTICE OF
THE POSSIBILITY OF SUCH DAMAGE.

1.F.3. LIMITED RIGHT OF REPLACEMENT OR REFUND - If

you discover a defect in this electronic work within 90 days of
receiving it, you can receive a refund of the money (if any) you
paid for it by sending a written explanation to the person you
received the work from. If you received the work on a physical
medium, you must return the medium with your written
explanation. The person or entity that provided you with the
defective work may elect to provide a replacement copy in lieu
of a refund. If you received the work electronically, the person or
entity providing it to you may choose to give you a second
opportunity to receive the work electronically in lieu of a refund.
If the second copy is also defective, you may demand a refund
in writing without further opportunities to fix the problem.

1.F.4. Except for the limited right of replacement or refund set

forth in paragraph 1.F.3, this work is provided to you ‘AS-IS’,
WITH NO OTHER WARRANTIES OF ANY KIND, EXPRESS
OR IMPLIED, INCLUDING BUT NOT LIMITED TO
WARRANTIES OF MERCHANTABILITY OR FITNESS FOR
ANY PURPOSE.

1.F.5. Some states do not allow disclaimers of certain implied

warranties or the exclusion or limitation of certain types of
damages. If any disclaimer or limitation set forth in this
agreement violates the law of the state applicable to this
agreement, the agreement shall be interpreted to make the
maximum disclaimer or limitation permitted by the applicable
state law. The invalidity or unenforceability of any provision of
this agreement shall not void the remaining provisions.

1.F.6. INDEMNITY - You agree to indemnify and hold the

Foundation, the trademark owner, any agent or employee of the
Foundation, anyone providing copies of Project Gutenberg™
electronic works in accordance with this agreement, and any
volunteers associated with the production, promotion and
distribution of Project Gutenberg™ electronic works, harmless
from all liability, costs and expenses, including legal fees, that
arise directly or indirectly from any of the following which you do
or cause to occur: (a) distribution of this or any Project
Gutenberg™ work, (b) alteration, modification, or additions or
deletions to any Project Gutenberg™ work, and (c) any Defect
you cause.

Section 2. Information about the Mission of

Project Gutenberg™
Project Gutenberg™ is synonymous with the free distribution of
electronic works in formats readable by the widest variety of
computers including obsolete, old, middle-aged and new
computers. It exists because of the efforts of hundreds of
volunteers and donations from people in all walks of life.

Volunteers and financial support to provide volunteers with the

assistance they need are critical to reaching Project
Gutenberg™’s goals and ensuring that the Project Gutenberg™
collection will remain freely available for generations to come. In
2001, the Project Gutenberg Literary Archive Foundation was
created to provide a secure and permanent future for Project
Gutenberg™ and future generations. To learn more about the
Project Gutenberg Literary Archive Foundation and how your
efforts and donations can help, see Sections 3 and 4 and the
Foundation information page at www.gutenberg.org.

Section 3. Information about the Project

Gutenberg Literary Archive Foundation
The Project Gutenberg Literary Archive Foundation is a non-
profit 501(c)(3) educational corporation organized under the
laws of the state of Mississippi and granted tax exempt status by
the Internal Revenue Service. The Foundation’s EIN or federal
tax identification number is 64-6221541. Contributions to the
Project Gutenberg Literary Archive Foundation are tax
deductible to the full extent permitted by U.S. federal laws and
your state’s laws.

The Foundation’s business office is located at 809 North 1500

West, Salt Lake City, UT 84116, (801) 596-1887. Email contact
links and up to date contact information can be found at the
Foundation’s website and official page at
www.gutenberg.org/contact

Section 4. Information about Donations to

the Project Gutenberg Literary Archive
Foundation
Project Gutenberg™ depends upon and cannot survive without
widespread public support and donations to carry out its mission
of increasing the number of public domain and licensed works
that can be freely distributed in machine-readable form
accessible by the widest array of equipment including outdated
equipment. Many small donations ($1 to $5,000) are particularly
important to maintaining tax exempt status with the IRS.

The Foundation is committed to complying with the laws

regulating charities and charitable donations in all 50 states of
the United States. Compliance requirements are not uniform
and it takes a considerable effort, much paperwork and many
fees to meet and keep up with these requirements. We do not
solicit donations in locations where we have not received written
confirmation of compliance. To SEND DONATIONS or
determine the status of compliance for any particular state visit
www.gutenberg.org/donate.

While we cannot and do not solicit contributions from states

where we have not met the solicitation requirements, we know
of no prohibition against accepting unsolicited donations from
donors in such states who approach us with offers to donate.

International donations are gratefully accepted, but we cannot

make any statements concerning tax treatment of donations
received from outside the United States. U.S. laws alone swamp
our small staff.

Please check the Project Gutenberg web pages for current

donation methods and addresses. Donations are accepted in a
number of other ways including checks, online payments and
credit card donations. To donate, please visit:
www.gutenberg.org/donate.

Section 5. General Information About Project

Gutenberg™ electronic works
Professor Michael S. Hart was the originator of the Project
Gutenberg™ concept of a library of electronic works that could
be freely shared with anyone. For forty years, he produced and
distributed Project Gutenberg™ eBooks with only a loose
network of volunteer support.

Project Gutenberg™ eBooks are often created from several

printed editions, all of which are confirmed as not protected by
copyright in the U.S. unless a copyright notice is included. Thus,
we do not necessarily keep eBooks in compliance with any
particular paper edition.

Most people start at our website which has the main PG search
facility: www.gutenberg.org.

This website includes information about Project Gutenberg™,

including how to make donations to the Project Gutenberg
Literary Archive Foundation, how to help produce our new
eBooks, and how to subscribe to our email newsletter to hear
about new eBooks.