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Animal Behavior

Eleventh edition

Dustin R. Rubenstein and John Alcock

Publication Date - June 2018

ISBN: 9781605355481

672 pages
9 x 10.875 inches

In Stock

Retail Price to Students: $129.99

Shows how researchers use scientific logic to study the underlying mechanisms and evolutionary bases of behavior, with emphasis on how evolutionary theory unifies the various subdisciplines within animal behavior.


A comparative and integrative overview of how and why animals as diverse as insects and humans behave the way that they do, linking behaviors to the brain, genes, and hormones, as well as to the surrounding ecological and social environments.

New to this Edition

  • New lead author, Dustin Rubenstein, widely known for his research and teaching accomplishments, is an integrative behavioral and evolutionary ecologist who studies the evolution of complex animal societies, and how organisms adapt to and cope with environmental change
  • A renewed focus on integration that links the evolution of behavior to neural, genetic, and physiological mechanisms
  • A return to its classic organizational structure, with proximate mechanisms introduced before an extended discussion of the ultimate factors underlying behavior, integrating both throughout
  • Introduction of cutting-edge neuro and genomic research approaches, all while maintaining a focus on the theoretical aspects of the field in an explicit hypothesis testing framework
  • Building on the foundation of previous editions, new pedagogical features designed to engage students and aid instructors, including Integrative Approaches, Exploring Behavior by Interpreting Data, Darwinian Puzzles, and Hypotheses Tables
  • QR codes provide students rapid access to actual audio clips of birdsongs, and high-definition video clips of animal behaviors
  • Updated and revised artwork that features more color, a more consistent palette for figures, crisper photographs, and error bars on figures


  • Engaging, narrative-driven text that draws from classic studies and primary literature
  • A classic approach to the evolutionary and comparative study of animal behavior, with a unique focus on integrative study
  • Taxonomically-diverse examples aid in understanding of the concepts, providing students with a clear idea of how scientists study behavior
  • Emphasis on hypothesis testing, observation and experimental methods, and discussion of costs and benefits
  • Extensive use of audio and visual collections, using QR codes for quick access

About the Author(s)

Dustin R. Rubenstein, Professor, Columbia University, USA

John Alcock, Professor Emeritus, Arizona State University, USA

Table of Contents

    Chapter 1. An Introduction to Animal Behavior
    Natural Selection and the Evolution of Behavior
    The Cost-Benefit Approach to Behavioral Biology
    The Levels of Analysis
    Box 1.1. Natural selection and infanticide in primates
    The Integrative Study of Animal Behavior
    Approaches to Studying Behavior
    The Adaptive Basis of Behavior: Mobbing in Gulls
    Box 1.2. Phylogenies and the comparative method
    Box 1.3. The benefit of high nest density for the arctic skua
    The Science of Animal Behavior
    The Integrative Study of Behavior

    Chapter 2. The Integrative Study of Behavior
    The Development of Song Learning
    Box 2.1. Characterizing sounds made by animals
    Intraspecific Variation and Dialects
    Social Experience and Song Development
    Mechanisms of Song Learning
    Box 2.2. Song learning in birds adoptedby another species
    The Genetics of Song Learning
    Control of the Avian Song System
    Box 2.3. Proximate mechanisms underlying song preferences in females
    The Evolution of Song Learning
    An Evolutionary History of Bird Song
    Mechanisms of Song Learning and the Comparative Approach
    Human versus Avian Vocal Learning
    The Adaptive Value of Song Learning
    Box 2.4. Why might song learning make males communicate more effectively with rivals or potential mates?
    Adapting to the Local Environment
    Recognition: Friends versus Foes
    Sexual Selection: Male-Male Competition
    Sexual Selection: Female Choice and Assortative Mating
    The Integrative Study of Bird Song

    Chapter 3. The Developmental and Genetic Bases of Behavior
    Behavior Requires Genes and the Environment
    The Interactive Theory of Development
    Box 3.1. Behavioral genetics: Identifying the genetic basis of differences in behavior
    Environmental Differences Can Cause Behavioral Differences
    Genetic Differences Can Also Cause Behavioral Differences
    Box 3.2. Migratory restlessness
    Learning and Cognition
    Learning Requires Both Genes and Environment
    Learning in Complex Environments
    The Adaptive Value of Learning
    The Evolutionary Development of Behavior
    The Evo-Devo Approach to Understanding Behavior
    Box 3.3. The genetics of foraging behavior in honey bees
    Early Life Developmental Conditions
    The Role of the Social Environment
    Developmental Homeostasis versus Developmental Constraint
    Developmental Switch Mechanisms
    Supergenes and Behavioral Polymorphisms

    Chapter 4. The Neural Basis of Behavior
    Responding to Stimuli
    Complex Responses to Simple Stimuli
    How Moths Avoid Bats
    Box 4.1. Ultrasound detection in the moth ear
    Ultrasonic Hearing in Other Insects
    Neural Command and Control
    Decision Making in the Brain
    From Ultrasound to Ultraviolet Radiation
    Selective Relaying of Sensory Inputs
    Responding to Relayed Messages
    The Proximate Basis of Stimulus Filtering
    Box 4.2. Determining how female parasitoid wasps choose their singing male bush-cricket hosts
    Box 4.3. Cortical magnification in mammals
    The Evolution of Cognitive Skills
    Box 4.4. Do energetic demands explain why humans have such large brains?

    Chapter 5. The Physiological Basis of Behavior
    Endogenous Rhythms and Changing Behavioral Priorities
    Mechanisms of Changing Behavioral Priorities
    The Neurobiology of Circadian Timing
    The Genetics of Circadian Timing
    The Physiology of Circadian Timing
    Seasonal and Annual Cycles of Behavior
    Cues That Entrain Cycles of Behavior
    Predictable Environmental Cues
    Box 5.1. Hormonal responses to light in birds
    Unpredictable Environmental Cues
    Social Conditions and Changing Priorities
    Hormonal Mechanisms Underlying Behavioral Change
    Organizational versus Activational Effects of Hormones on Behavior and Development
    Box 5.2. Measuring hormones in animals
    Box 5.3. Do steroid hormones modulate male parental behavior in California mice?
    Hormones and Reproduction
    Testosterone and Reproductive Behavior
    The Costs of Hormonal Regulation
    Glucocorticoids and Responding to Environmental Change

    Chapter 6. Avoiding Predators and Finding Food
    Avoiding Predators
    Social Defenses
    Box 6.1. Evolutionary game theory
    Game Theory and Social Defenses
    Box 6.2. Game theory and the selfish herd
    Blending In
    Standing Out
    Optimality Theory and Antipredator Behavior
    Finding Food
    Optimality Theory and Foraging Decisions
    Box 6.3. Territoriality and feeding behavior in golden-winged sunbirds
    Box 6.4. Optimal foraging by pike cichlid fish
    Criticisms of Optimal Foraging Theory
    Landscapes of Fear
    Game Theory and Feeding Behavior

    Chapter 7. Territoriality and Migration
    Where to Live
    Habitat Selection
    Territoriality and Resource-Holding Potential
    Box 7.1. How to track migratory songbirds
    Why Give Up Quickly When Fighting for a Territory?
    The Dear Enemy Effect
    To Stay or Go
    Box 7.2. Opposite patterns of sex-biased dispersal in mammals and birds
    The Costs and Benefits of Migration
    Box 7.3. Behaviors to reduce the costs of flying during migration
    Variation in Migratory Behavior
    Box 7.4. Migratory pathways of Swainson's thrush

    Chapter 8. Principles of Communication
    Communication and Animal Signals
    Information Use and Animal Signals
    The Evolution of Animal Signals
    Preexisting Traits and the Development of a Strange Display
    The Panda Principle and Preexisting Traits
    Preexisting Biases and the Evolution of Animal Signals
    Box 8.1. Spiders hunting prey at night
    Preexisting Traits versus Preexisting Biases
    Box 8.2. Why do female moths mate with males that produce ultrasonic mimetic signals similar to those produced by predatory bats?
    The Function of Animal Signals
    The Adaptive Function of a Strange Display
    Honest Communication and Threat Displays
    Honest Signaling
    Box 8.3. Mechanisms and measurement of animal coloration
    When Multiple Honest Signals Are Better Than One
    Deceitful Signaling
    Eavesdropping on Others

    Chapter 9. Reproductive Behavior
    Sexual Selection and the Evolution of Sex Differences
    Sex Differences in Reproductive Behavior
    Box 9.1. Are sperm always cheap?
    Sex Differences and Parental Investment
    A Reversal in Sex Differences
    Intrasexual Selection and Competition for Mates
    Competition and Access to Mates
    Coexistence of Conditional Mating Tactics
    Coexistence of Alternative Mating Strategies
    Sperm Competition
    Mate Guarding and Paternity Assurance
    Intersexual Selection and Mate Choice
    Female Mate Choice for Direct Benefits
    Female Mate Choice for Indirect Benefits
    Box 9.2. Sexual selection in the peacock
    Runaway versus Chase-away Sexual Selection
    Cryptic Female Choice
    Sexual Conflict
    The Manipulation of Female Choice
    Sexual Arms Races

    Chapter 10. Mating Systems
    Monogamy: A Lack of Multiple Mating
    Why Be Monogamous?
    Monogamy in Species with Paternal Care
    Monogamy When Paternal Care Is Rare
    Polyandry: Multiple Mating by Females
    Monogamous Males and Polyandrous Females
    Box 10.1. Sexual parasitism, dwarf males, and the evolution of gigolos
    Polyandry and Indirect Genetic Benefits
    Box 10.2. Extra-pair paternity and good genes in birds
    Polyandry and Direct Benefits
    Polygyny: Multiple Mating by Males
    Female Defense Polygyny
    Resource Defense Polygyny
    Lek Polygyny
    Scramble Competition Polygyny
    Box 10.3. Lekking females in a sex-role reversed pipefish
    Polygynandry and Promiscuity: Multiple Mating by Both Sexes

    Chapter 11. Parental Care
    Offspring Value and Parental Investment
    Parental Care Decisions
    Parental Favoritism in Offspring Care and Production
    Parental Favoritism in Humans
    Family Conflict
    To Care or Not to Care
    The Costs and Benefits of Parental Care
    Sexual Conflict and Parental Care: Who Cares?
    Box 11.1. Why do females provide all of the care in treehoppers?
    Why Do Females Care?
    Why Do Males Care?
    Box 11.2. Reactions of nest-defending bluegill males to potential egg and fry predators under two conditions
    Discriminating Parental Care
    Recognizing One's Own Offspring
    Box 11.3. Why do parents in some species adopt genetic strangers of their own species?
    Interspecific Brood Parasitism
    Choosing the Correct Host
    Coevolutionary Arms Races
    The Evolution of Interspecific Brood Parasitism

    Chapter 12. Principles of Social Evolution
    Box 12.1. The major evolutionary transitions
    Altruism and the Levels of Selection
    Individual versus Group Selection
    Altruism and the Role of Kin Selection
    Box 12.2. Calculating genetic relatedness
    Kin Selection and Inclusive Fitness Theory
    Challenges to Kin Selection and Inclusive Fitness Theory
    Box 12.3. Altruism in amoebae
    Haplodiploidy and the Evolution of Eusociality
    Testing the Haplodiploidy Hypothesis
    Inclusive Fitness and Monogamy in Eusocial Insects
    Box 12.4. Division of labor in clonal trematode flatworms
    Sterility and Caste Differentiation
    Social Conflict in Animal Societies
    Reproductive Conflict

    Chapter 13. Social Behavior and Sociality
    The Evolution of Social Behavior
    Forms of Social Behavior
    Mutual Benefit
    Box 13.1. How do groups of animals decide where to go?
    Box 13.2. Social network analysis
    Altruism and Reciprocity
    Non-cooperative Social Behaviors: Selfishness and Spite
    Individual Differences in Social Behavior
    Personalities in Social Species
    The Evolution of Cooperative Breeding
    Reproductive Cooperation and Kin Selection
    Reproductive Benefits and Cooperative Breeding
    Box 13.3. Mobbing and kinship in groups of Siberian jays
    Reproductive Costs and Cooperative Breeding
    Reproductive Conflict in Cooperative Breeders
    Reproductive Suppression
    Reproductive Skew, Extra-pair Paternity, and Social Structure
    Box 13.4. Why do males and females both have elaborate traits in social species?

    Chapter 14. Human Behavior
    The Development and Evolutionary History of Human Speech
    Box 14.1. Ethical studies of humans and other animals
    The Neurophysiology of Speech
    The Adaptive Value of Speech
    Reproductive Behavior
    An Evolutionary Analysis of Human Mate Choice
    Mate Choice by Women
    Box 14.2. Female choice and the features of dominant versus attractive men
    Box 14.3. Human mate choice in an online world
    Mate Choice by Men
    Reproductive and Sexual Conflict in Humans
    Extreme Sexual Conflict in Humans: Polygamy and Extramarital Affairs
    Coercive Sex
    Practical Applications of Behavioral Theory
    Evolutionary Medicine
    The Triumph of an Evolutionary Analysis of Human Behavior

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