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Principles of Animal Behavior, 4th Edition

Fourth Edition

Since the last edition of this definitive textbook was published in 2013, much has happened in the field of animal behavior. In this fourth edition, Lee Alan Dugatkin draws on cutting-edge new work not only to update and expand on the studies presented, but also to reinforce the previous editions’ focus on ultimate and proximate causation, as well as the book’s unique emphasis on natural selection, learning, and cultural transmission. The result is a state-of-the-art textbook on animal behavior that explains underlying concepts in a way that is both scientifically rigorous and accessible to students. Each chapter in the book provides a sound theoretical and conceptual basis upon which the empirical studies rest. A completely new feature in this edition are the Cognitive Connection boxes in Chapters 2–17, designed to dig deep into the importance of the cognitive underpinnings to many types of behaviors. Each box focuses on a specific issue related to cognition and the particular topic covered in that chapter.

As Principles of Animal Behavior makes clear, the tapestry of animal behavior is created from weaving all of these components into a beautiful whole. With Dugatkin’s exquisitely illustrated, comprehensive, and up-to-date fourth edition, we are able to admire that beauty anew.

See resources for instructors.
Read the first chapter.

576 pages | 529 color plates, 31 halftones, 3 line drawings | 8 1/2 x 11 | © 2020

Biological Sciences: Anatomy, Behavioral Biology, Evolutionary Biology, Physiology, Biomechanics, and Morphology


“Up-to-date, highly integrative, and richly illustrated. It thus merits serious consideration by anyone looking for a textbook to support undergraduate offerings in animal behavior or behavioral ecology. . . . Principles of Animal Behavior is comprehensive and readable, summarizing not only what is well documented but more importantly where integrated understanding is lacking, and thus where further research will prove most profitable.”

Animal Behaviour, on the first edition

“The book reveals a richly illustrated panoramic view of animal behavior and, where it can, it also provides examples of the physiological, neurobiological, and molecular genetic mechanisms that may underlie it. . . . Dugatkin’s text . . . can be enjoyed by anyone who has an interest in the beauty of animal behavior. . . . Excellent.”

Times Higher Education, on the second edition

"Dugatkin offers a highly readable overview of the modern field of animal behavior, keeping up with the exponential growth over the last few decades. Instead of the usual choice between inborn and learned behavior, this book offers the reader with a fully integrated view."

Frans de Waal, Emory University | on the third edition

"This is an up-to-date text that shows not only how theory and empirical data are combined, but how they are derived on the ground."

Bernd Heinrich, University of Vermont | on the third edition

Table of Contents

1 Principles of Animal Behavior
Types of Questions and Levels of Analysis
What Is Behavior?
Three Foundations
Natural Selection
Individual Learning
Cultural Transmission
Conceptual, Theoretical, and Empirical Approaches
Conceptual Approaches
Theoretical Approaches
Empirical Approaches
An Overview of What Is to Follow
Interview with Dr. E. O. Wilson

2 The Evolution of Behavior
Artificial Selection
Natural Selection
Selective Advantage of a Trait
How Natural Selection Operates
Sociobiology, Selfish Genes, and Adaptation
CONSERVATION CONNECTION: Conservation Biology and Symmetry as an Indicator of Risk
Antipredator Behavior in Guppies
Kinship and Naked Mole Rat Behavior
COGNITIVE CONNECTION: Tool Use in New Caledonian Crows
Phylogeny and the Study of Animal Behavior
Phylogenetic Trees
Phylogeny and Parental Care
Phylogeny and Courtship Behavior
Interview with Dr. Alan Grafen

3 Hormones and Neurobiology
Ultimate and Proximate Perspectives
Proximate Causation: Hormones
CONSERVATION CONNECTION: Community-Based Ecotourism: Using Hormones to Measure Effects on Animal Well-Being
How the Endocrine System Integrates Sensory Input and Output
The Long-Term Effects of In Utero Exposure to Hormones
Vasopressin and Sociality in Voles
Hormones and Honeybee Foraging
Hormones and Vocalizations in Plainfin Midshipman Fish
Neurobiological Underpinnings of Behavior
The Nervous Impulse
Mushroom Bodies and Honeybee Foraging
COGNITIVE CONNECTION: Brain Size and Problem Solving
Neurobiology and Vocalizations in Plainfin Midshipman Fish
Sleep and Predation in Mallard Ducks
Interview with Dr. Geoffrey Hill

4 Molecular Genetics and Development
Molecular Genetics and Animal Behavior
Mendel’s Laws
Locating Genes for Polygenic Traits
Genes, mRNA, and Honeybee Foraging
Song Acquisition in Birds
avpr1a, Vasopressin, and Sociality in Voles
Genetic Toolkits, Transcription Factors, and Territoriality
Development and Animal Behavior
Development, Temperature, and Ovipositing Behavior in Wasps
CONSERVATION CONNECTION: Development, Dispersal, and Climate Change
Family Structure, Development, and Behavior in Prairie Voles
Early Nest Development and Behavior in Cichlid Fish
Early Development and Its Effect on Parental Behavior in the Oldfield Mouse
Interview with Dr. Gene Robinson

5 Learning
What Is Individual Learning?
How Animals Learn
Learning from a Single-Stimulus Experience
Pavlovian (Classical) Conditioning
Instrumental (Operant) Conditioning
Why Animals Learn
Within-Species Studies and the Evolution of Learning
COGNITIVE CONNECTION: Natural Selection and Associative Learning
Population Comparisons and the Evolution of Learning
A Model of the Evolution of Learning
What Animals Learn
Learning about Predators
CONSERVATION CONNECTION: Learning, Alarm Chemicals, and Reintroduction Programs
Learning about Their Mate
Learning about Familial Relationships
Learning about Aggression
Molecular Genetics and Endocrinology of Learning
Molecular Genetics of Learning in Rats
Endocrinology of Learning in Rats
Interview with Dr. Sara Shettleworth
6 Cultural Transmission
What Is Cultural Transmission?
What’s So Important about Cultural Transmission?
Effects of Others on Behavior
Social Learning
CONSERVATION CONNECTION: Crop Raiding, Elephants, and Social Learning
The Rise and Fall of a Tradition
Teaching in Animals
COGNITIVE CONNECTION: Parents Teaching Embryos?
Modes of Cultural Transmission
Vertical Cultural Transmission
Oblique Cultural Transmission
Horizontal Cultural Transmission
The Interaction of Genetic and Cultural Transmission
The Grants’ Finches
Guppy Mate Choice
Cultural Transmission and Brain Size
Interview with Dr. Cecilia Heyes

7 Sexual Selection
COGNITIVE CONNECTION: Aggression, Observation, and Gene Expression in the Brain of Females
Intersexual and Intrasexual Selection
Evolutionary Models of Mate Choice
CONSERVATION CONNECTION: Genetic Diversity, Genetic Quality, and Conservation Biology
Direct Benefits and Mate Choice
Good Genes and Mate Choice
Runaway Sexual Selection
Sensory Bias and the Emergence of Mate Choice
Learning and Mate Choice
Sexual Imprinting
Learning and Mate Choice in Japanese Quail
Cultural Transmission and Mate Choice
Mate-Choice Copying
Song Learning and Mate Choice in Cowbirds
Male-Male Competition and Sexual Selection
Red Deer Roars and Male-Male Competition
Male-Male Competition by Interference
Male-Male Competition by Cuckoldry
Interview with Dr. Anne Houde

8 Mating Systems
Different Mating Systems
Monogamous Mating Systems
Polygamous Mating Systems
COGNITIVE CONNECTION: The Social Brain Hypothesis
Promiscuous Mating Systems
The Ecology and Evolution of Polygynous Mating Systems
Polygyny and Resources
The Polygyny Threshold Model
CONSERVATION CONNECTION: Anthropogenic Effects on Animal Mating Systems
Extrapair Copulations
Sperm Competition
Sperm Cooperation
Multiple Mating Systems in a Single Population?
Interview with Dr. Catherine Marler

9 Kinship
Kinship and Animal Behavior
Kinship Theory
Relatedness and Inclusive Fitness
Family Dynamics
CONSERVATION CONNECTION: Nonbreeding Groups and Inclusive Fitness Benefits in Gorillas
Conflict within Families
Parent-Offspring Conflict
Sibling Rivalry
Kin Recognition
Matching Models
COGNITIVE CONNECTION: Social Learning, Kinship, and Antipredator Behavior
Interview with Dr. Francis Ratnieks

10 Cooperation
Defining Cooperation
Paths to Cooperation
Path 1: Reciprocity
Path 2: Byproduct Mutualism
Path 3: Group Selection
CONSERVATION CONNECTION: Cooperation, the Tragedy of the Commons, and Overharvesting
Coalitions in Baboons
Alliances and “Herding” Behavior in Cetaceans
A Phylogenetic Approach to Cooperation
Phylogeny and Cooperative Breeding in Birds
Phylogeny and Cooperation in Shrimp
Phylogeny and Cooperation in Social Spiders
Interspecific Mutualisms
Ants and Butterflies—Mutualism with Communication?
Coral and Coral Reef Fish
Interview with Dr. Hudson Kern Reeve

11 Foraging
Finding Food and the Search Image
Optimal Foraging Theory
What to Eat
Where to Eat
Risk-Sensitive Foraging
Growing Food
Foraging and Group Life
Group Size
Groups, Public Information, and Foraging
Natural Selection, Phylogeny, and Seed Caching
Hippocampal Size and Caching Ability
A Phylogenetic Approach to Studying Caching Ability
Learning and Foraging
COGNITIVE CONNECTION: Proximate Factors in Foraging-Related Learning
Foraging, Learning, and Brain Size in Birds
CONSERVATION CONNECTION: Behavioral Traditions, Foraging, and Conservation in Killer Whales
Foraging Innovation and Diversification in Emberizoidea
Social Learning and Foraging
Interview with Dr. John Krebs

12 Antipredator Behavior
Avoiding Predators
Blending into the Environment
Being Quiet
Choosing Safe Habitats
CONSERVATION CONNECTION: Co-evolution, Naive Prey, and Introduction Programs
What Prey Do When They Encounter Predators
COGNITIVE CONNECTION: Heritability of Conditioned Fear Responses
Approaching Predators
Feigning Death
Signaling to Predators
Fighting Back
Predation and Foraging Trade-offs
Interview with Dr. Anne Magurran

13 Communication
Communication and Honesty
Communication Solves Problems
Problem: How to Coordinate Group Foraging
Problem: How to Find and Secure a Mate
CONSERVATION CONNECTION: Anthropogenic Change and Animal Communication
Problem: Predators
COGNITIVE CONNECTION: Can Elephants Distinguish Between Humans Based on Voice?
Interview with Dr. Rufus Johnstone

14 Habitat Selection, Territoriality, and Migration
Habitat Choice
The Ideal Free Distribution Model and Habitat Choice
Avoidance of Disease-Filled Habitats
Stress Hormones, Spatial Memory, and Habitat Choice in Rats
Territoriality and Learning
COGNITIVE CONNECTION: Nest Complexity and Cerebellar Foliation
Conflict within Family Territories
Multiple Migratory Routes
Migration and Navigation
CONSERVATION CONNECTION: Migration Patterns, “Stopovers,” and Conservation Biology
Migration, Temperature, and Basal Metabolic Rate
Migration and Defense against Parasites
A Phylogenetic Approach to Migratory Behavior
Interview with Dr. Judy Stamps

15 Aggression
Fight or Flight?
CONSERVATION CONNECTION: Breeding Programs Can Lead to More Aggressive Animals
Game Theory Models of Aggression
The Hawk-Dove Game
The War of Attrition Model
The Sequential Assessment Model
Winner, Loser, Bystander, and Audience Effects
Winner and Loser Effects
Bystander Effects
Audience Effects
Aggression and Social Network Theory
COGNITIVE CONNECTION: Aggression, Observation, and Gene Expression in the Brain of Males
Interview with Dr. Karen Hollis

16 Play
Defining Play
CONSERVATION CONNECTION: Play Behavior as a Measure of Environmental Stress
Types and Functions of Play
Object Play
Locomotor Play
Social Play
COGNITIVE CONNECTION: Play and Brain Development
A General Theory for the Function of Play
Endocrinological and Neurobiological Bases of Play
Play Fighting in Young Male Rodents
A Phylogenetic Approach to Play
Interview with Dr. Marc Bekoff

17 Animal Personalities
Boldness and Shyness
Bold and Shy Pumpkinseeds
Some Case Studies
Hyena Personalities
Octopus and Squid Personalities
Natural Selection and Personality in Great Tits
Coping Styles
COGNITIVE CONNECTION: Brain Size and the Proactive-Reactive Personality Continuum
CONSERVATION CONNECTION: Using Personality to Reduce Human-Animal Conflicts
Personality and Dispersal Behavior
Interview with Dr. Sam Gosling


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