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Animal Behavior

Concepts, Methods, and Applications

Third Edition

Shawn E. Nordell and Thomas J. Valone

Publication Date - July 2020

ISBN: 9780190924256

560 pages

In Stock

Retail Price to Students: $97.99

A conceptual approach that puts the process of science and applications front and center


EMPHASIZES CONCEPTS. Animal Behavior: Concepts, Methods, and Applications, Third Edition, uses broad organizing concepts to provide a framework for understanding the science of animal behavior. In an engaging, question-driven style, Shawn E. Nordell and Thomas J. Valone offer readers a clear learning progression for understanding and evaluating empirical research examples.

FOCUSES ON METHODOLOGY AND THE PROCESS OF SCIENCE. Featured studies illustrate each concept and emphasize the experimental designs and the hypothesis testing methods scientists use to address research questions.

HIGHLIGHTS REAL-WORLD APPLICATIONS. Concepts are related to real life to help students understand the broader significance of animal behavior research, including applications to human behavior and conservation.

New to this Edition

  • Carefully crafted Learning Objectives help students guide their learning and measure intended learning outcomes of the material
  • A Chapter Reviews provide a brief summary of major points of each concept
  • A new Chapter 16: Cooperative Behavior includes expanded coverage of inclusive fitness theory and kin discrimination; additional coverage of game theory, including the prisoner's dilemma and snowdrift games; and expanded coverage of the evolution of cooperative reproduction
  • New Quantitative Reasoning (online) exercises help students develop their data and graphical analysis skills
  • New Toolbox on Interpreting Graphical Data will help students develop the quantitative skills of reading the data, qualitative skills of reading between the data, and analytical skills of reading beyond the data
  • Incorporates recent research developments, including genomics
  • Additional coverage of proximate explanations with expanded discussion of Tinbergen's 4 Questions
  • Expands and revises the Video Guide and Test Bank
  • Includes Critical Thinking and Discussion Questions to foster in-class discussions
  • A new dynamic enhanced eBook brings the study of animal behavior to life as never before, engaging students through embedded and linked videos as well as interactive Quantitative Reasoning Exercises


  • NEW! Learning Objectives: Each concept includes objectives to help students guide and assess their learning of the material
  • NEW! Flashcards: Key Terms are provided in flashcard format for student self-testing
  • NEW! Chapter Review: Each chapter now includes a review that summarizes the major points of each concept
  • NEW! Quantitative Reasoning (online) exercises: These exercises help students develop their data and graphical analysis skills. They present students with data from a research study and ask questions that require them to extract specific information from the data, understand the relationships among variables, and make inferences and predictions from data.
  • "Scientific Process" boxes clearly and concisely lay out each step of the scientific process for highlighted research studies
  • "Toolboxes" provide in-depth coverage of mathematical models and essential behavior research skills
  • "Applying the Concepts" boxes provide examples of how animal behavior research is being applied to real-life problems
  • Diverse research examples represent a broad array of taxa from around the world and highlight both classic and contemporary research from a diverse set of researchers
  • Beautiful illustrations and photos help readers visualize complex processes and connect with the natural world

About the Author(s)

Shawn E. Nordell is Educational and Career Specialist at Washington University in St. Louis.

Thomas J. Valone is Professor of Biology and Director of the Reis Biological Station at Saint Louis University.

Table of Contents


    Chapter 1. The Science of Animal Behavior
    1.1 Animals and their behavior are an integral part of human society
    Recognizing and defining behavior
    Measuring behavior in elephant ethograms
    1.2 The scientific method is a formalized way of knowing about the natural world
    The importance of hypotheses
    The scientific method
    Negative results and directional hypotheses
    Correlation and causality
    Hypotheses and theories
    Social sciences and the natural sciences
    1.3 Scientists study both the proximate mechanisms that generate behavior and the ultimate reasons why the behavior evolved
    Tinbergen's four questions
    Implications of Tinbergen's work
    1.4 Researchers have examined animal behavior from a variety of perspectives over time
    Darwin and adaptation
    Early comparative psychology
    Comparative psychology in North America
    Classical ethology
    Interdisciplinary approaches
    1.5 Anthropomorphic explanations of behavior assign human emotions to animals and can be difficult to test
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 1.1 Robin abundance and food availability
    Scientific Process 1.2 Robin abundance and predators
    Applying the Concepts 1.1 Human infant crying
    Applying the Concepts 1.2 What is behind the "guilty look" in dogs?
    Toolbox 1.1 Describing and summarizing data
    Toolbox 1.2 Interpreting graphical data

    Quantitative Reasoning 1.1 Nesting success and breeding habitats

    Chapter 2. Methods for Studying Animal Behavior
    2.1 Animal behavior scientists generate and test hypotheses to answer research questions about behavior
    Hypothesis testing in wolf spiders
    Generating hypotheses
    Hypotheses and predictions from mathematical models
    2.2 Researchers use observational, experimental, and comparative methods to study behavior
    The observational method
    The observational method and male mating tactics in bighorn sheep
    The experimental method
    The experimental method and jumping tadpoles
    The comparative method
    The comparative method and the evolution of burrowing behavior in mice
    2.3 Animal behavior research requires ethical animal use
    How research can affect animals
    Sources of ethical standards
    The three Rs
    2.4 Scientific knowledge is generated and communicated to the scientific community via peer-reviewed research
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 2.1 Jumping tadpoles
    Applying the Concepts 2.1 Project Seahorse
    Toolbox 2.1 Animal sampling techniques
    Toolbox 2.2 Scientific literacy

    Quantitative Reasoning 2.1 Sampling methods

    Chapter 3. Evolution and the Study of Animal Behavior
    3.1 Evolution by natural selection favors behavioral adaptations that enhance fitness
    Measures of heritability
    Maternal defense behavior in mice
    Variation within a population
    Frequency-dependent selection
    Fitness and adaptation
    3.2 Modes of natural selection describe population changes
    Directional selection in juvenile ornate tree lizards
    Disruptive selection in spadefoot toad tadpoles
    Stabilizing selection in juvenile convict cichlids
    Studying adaptation: the cost-benefit approach
    3.3 Individual and group selection have been used to explain cooperation
    3.4 Sexual selection is a form of natural selection that focuses on the reproductive fitness of individuals
    Sexual selection in widowbirds
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 3.1 Stabilizing selection on territory size in cichlids
    Applying the Concepts 3.1 Do lemmings commit suicide?
    Toolbox 3.1 Genetics primer

    Quantitative Reasoning 3.1 Presence and absence of predator cues

    Chapter 4. Behavioral Genetics
    4.1 Behaviors vary in their heritability
    4.2 Behavioral variation is associated with genetic variation
    Behavioral differences between wild-type and mutant-type fruit flies
    Major and minor genes
    Fire ant genotype and social organization
    Experimental manipulation of gene function: knockout studies
    Anxiety-related behavior and knockout of a hormone receptor in mice
    QTL mapping to identify genes associated with behavior
    QTL mapping for aphid feeding behavior
    4.3 The environment influences behavior via gene expression
    Environmental effects on zebrafish aggression
    Social environment and gene expression in fruit flies
    Social environment and birdsong development
    Social environment and gene expression in birds
    Gene-environment interactions
    Rover and sitter foraging behavior in fruit flies
    4.4 Genomic approaches correlate gene expression with behavioral phenotypes
    Scouting behavior in bees
    Genomics and alternative mating tactics in fish
    4.5 Genes can limit behavioral flexibility
    Bold and shy personalities in streamside salamanders
    Aggressive personalities in funnel-web spiders
    Animal personalities model with fitness trade-offs
    Environmental effects on jumping spider personalities
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 4.1 Environmental effects on zebrafish aggression
    Scientific Process 4.2 Heritability of great tit exploratory behavior
    Scientific Process 4.3 Salamander personalities
    Applying the Concepts 4.1 Dog behavior heritability
    Toolbox 4.1 Molecular techniques

    Quantitative Reasoning 4.1 Female body size and sexual cannibalism

    Chapter 5. Sensory Systems and Behavior
    5.1 Animals acquire environmental information from their sensory systems
    5.2 Chemosensory systems detect chemicals that are perceived as tastes and odors
    Sweet and umami taste perception in rodents
    Cuttlefish physiological response to odors
    5.3 Photoreception allows animals to detect light and perceive objects as images
    Color vision in monarch butterflies
    Ultraviolet plumage reflectance in birds
    Infrared detection in snakes
    5.4 Mechanoreceptors detect vibrations that travel through air, water, or substrates
    Ultrasonic song detection in moths
    Long-distance communication in elephants
    Catfish track the wake of their prey
    Substrate-borne vibrations
    Antlions detect substrate-borne vibrations
    5.5 Some animals can detect electric or magnetic fields
    Sharks detect electric fields
    5.6 Predator and prey sensory systems co-evolve
    Insect tympanal organs: an evolved antipredator adaptation
    Predator-prey sensory system co-evolution in bats and moths
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 5.1 Antlion mechanoreception
    Applying the Concepts 5.1 How do mosquitoes find their victims?

    Quantitative Reasoning 5.1 Hummingbird hawkmoths and sugar preference

    Chapter 6. Communication
    6.1 Communication occurs when a specialized signal from one individual influences the behavior of another
    Honeybees and the waggle dance
    Odor or the waggle dance in bees
    Auditory signals: alarm calls
    Titmouse alarm calls
    Information or influence?
    6.2 The environment influences the evolution of signals
    Temperature affects ant chemical signals
    Habitat light environment affects fish visual signals
    Habitat structure affects bowerbird auditory signals
    6.3 Signals often accurately indicate signaler phenotype and environmental conditions
    Signals as accurate indicators: theory
    Aposematic coloration in frogs
    Courtship signaling in spiders
    Aggressive display and male condition in fighting fish
    6.4 Signals can be inaccurate indicators when the fitness interests of signaler and receiver differ
    Batesian mimicry and Enstaina salamanders
    Aggressive mimicry in fangblenny fish
    Intraspecific deception: false alarm calls
    Topi antelope false alarm calls
    Capuchin monkeys and inaccurate signals
    6.5 Communication can involve extended phenotype signals
    Bowerbirds construct and decorate bowers
    Sticklebacks decorate their nests
    6.6 Communication networks affect signaler and receiver behavior
    Squirrel eavesdropping
    Audience effects in fighting fish
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 6.1 Signaling in male wolf spiders
    Scientific Process 6.2 Fighting fish opercular display
    Applying the Concepts 6.1 Pheromones and pest control
    Applying the Concepts 6.2 Urban sounds affect signal production
    Applying the Concepts 6.3 Human luxury brands as costly signals

    Quantitative Reasoning 6.1 Sand hoods as extended phenotype signals

    Chapter 7. Learning, Neuroethology, and Cognition
    7.1 Learning allows animals to adapt to their environment
    Improved foraging efficiency in salamanders
    Evolution of learning
    Fiddler crab habituation
    7.2 Learning is associated with neurological changes
    Neurotransmitters and learning in chicks
    Dendritic spines and learning in mice
    Avian memory of stored food
    7.3 Animals learn associations between stimuli and responses
    Classical conditioning
    Pavlovian conditioning for mating opportunities in Japanese quail
    Fish learn novel predators
    Operant conditioning
    Learning curves in macaques
    Trial-and-error learning in bees
    7.4 Social interactions facilitate learning
    Learned anti-predator behaviors in prairie dogs
    Learning about food patches
    Social information use in sticklebacks
    Ptarmigan hens teach chicks their diet
    Tandem running in ants
    7.5 Social learning can lead to the development of animal traditions and culture
    Foraging behavioral traditions in great tits
    7.6 Animals vary in their cognitive abilities
    Tool use in capuchin monkeys
    Problem solving and insight learning
    Insight learning in keas
    Numerical competency in New Zealand robins
    Cognition and brain architecture in birds
    Brain size and cognition in guppies
    Cognitive performance and fitness in bowerbirds
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 7.1 Brain structure and food hoarding
    Scientific Process 7.2 Fish learn predators
    Applying the Concepts 7.1 Operation Migration and imprinting
    Applying the Concepts 7.2 Dog training
    Applying the Concepts 7.3 Human social learning about dangerous animals

    Quantitative Reasoning 7.1 Aggressiveness and learning ability

    Chapter 8. Foraging Behavior
    8.1 Animals find food using a variety of sensory modalities
    Bees use multiple senses to enhance foraging efficiency
    Gray mouse lemurs use multiple senses to find food
    8.2 Visual predators find cryptic prey more effectively by learning a search image
    Trout and search images
    8.3 The optimal diet model predicts the food types an animal should include in its diet
    The diet model
    A graphical solution
    Diet choice in northwestern crows
    Ant foraging and the effect of nutrients
    8.4 The optimal patch-use model predicts how long a forager should exploit a food patch
    The optimal patch-use model
    Patch use by ruddy ducks
    Optimal patch model with multiple costs
    Fruit bats foraging on heterogeneous patches
    Kangaroo rat foraging with variable predation costs
    Incomplete information and food patch estimation
    Bayesian foraging bumblebees
    8.5 Some animals obtain food from the discoveries of others
    Spice finch producer-scrounger game
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 8.1 Prey detection by gray mouse lemurs
    Scientific Process 8.2 Cryptic prey reduces predator efficiency
    Scientific Process 8.3 Patch use by fruit bats
    Applying the Concepts 8.1 Human patch-leaving decisions
    Applying the Concepts 8.2 GUDs and conservation
    Toolbox 8.1 Mathematical solution to the optimal diet model

    Quantitative Reasoning 8.1 Foraging in different habitats

    Chapter 9. Antipredator Behavior
    9.1 Animals reduce predation risk by avoiding detection
    Predator avoidance by cryptic coloration in crabs
    Predators and reduced activity in lizards
    Prey take evasive or aggressive action when detected
    Startle display in butterflies
    9.2 Many behaviors represent adaptive trade-offs involving predation risk
    Increased vigilance decreases feeding time
    Vigilance and predation risk in elk
    Rich but risky
    Environmental conditions and predation risk in foraging redshanks
    Mating and refuge use in fiddler crabs
    Perceived predation risk affects reproductive behavior in sparrows
    9.3 Living in groups can reduce predation risk
    The dilution effect and killifish
    The selfish herd and vigilance behavior
    Group size effect and the selfish herd hypothesis in doves
    9.4 Some animals interact with predators to deter attack
    Predator harassment in ground squirrels
    Pursuit deterrence and alarm signal hypotheses
    Tail-flagging behavior in deer
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 9.1 Feeding trade-off in redshanks
    Scientific Process 9.2 Predator harassment by California ground squirrels
    Applying the Concepts 9.1 Human fear of predators
    Applying the Concepts 9.2 Mitigating crop damage by manipulating predation risk

    Quantitative Reasoning 9.1 Anti-predator vigilance in yellow-bellied marmots

    Chapter 10. Dispersal and Migration
    10.1 Dispersal reduces resource competition and inbreeding
    Density-dependent dispersal in earthworms
    Food-related dispersal in water boatmen
    Inbreeding avoidance in great tits
    10.2 Reproductive success and public information affect breeding dispersal behavior
    Reproductive success and breeding dispersal in dragonflies
    Public information and breeding dispersal in kittiwakes
    10.3 Individuals migrate in response to changes in the environment
    Migration and changing resources
    Resource variation and migration in neotropical birds
    Heritability of migration behavior in Eurasian blackcaps
    A model of the evolution of migration
    Competition and migratory behavior of newts
    Maintenance of polymorphism in migratory behavior
    Alternative migratory behaviors in dippers
    10.4 Environmental cues and compass systems are used for orientation when migrating
    Compass systems
    Antennae and the sun compass system in monarchs
    The magnetic compass in sea turtles
    Multimodal orientation
    10.5 Bicoordinate navigation allows individuals to identify their location relative to a goal
    Bicoordinate navigation and magnetic maps in sea turtles
    Bicoordinate navigation in birds
    Homing migration in salmon
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 10.1 Breeding dispersal in dragonflies
    Scientific Process 10.2 The role of the antennae in the monarch butterfly sun compass
    Applying the Concepts 10.1 Bird migration and global climate change
    Applying the Concepts 10.2 Citizen scientists track fall migration flyways of monarch butterflies
    Applying the Concepts 10.3 Human magnetic orientation
    Toolbox 10.1 Emlen funnels

    Quantitative Reasoning 10.1 Dispersing cane toads

    Chapter 11. Habitat Selection, Territoriality, and Aggression
    11.1 Resource availability and the presence of others can influence habitat selection
    The ideal free distribution model
    The ideal free distribution model and guppies
    The ideal free distribution model and pike
    Cuckoos assess habitat quality
    Conspecific attraction
    Conspecific attraction and Allee effects in grasshoppers
    Conspecific cueing in American redstarts
    11.2 Individual condition and environmental factors affect territoriality
    Body condition and territoriality in damselflies
    Environmental factors and territory size in parrotfish
    11.3 Hormones influence aggression
    Winner-challenge effect in the California mouse
    Challenge hypothesis and bystanders in fish
    Juvenile hormone and wasp aggression
    11.4 Game theory models explain how the decisions of opponents and resource value affect fighting behavior
    The hawk-dove model
    Wrestling behavior in red-spotted newts
    Game theory assessment models
    Fiddler crab contests over burrows
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 11.1 Ideal free guppies
    Scientific Process 11.2 Conspecific attraction in grasshoppers
    Applying the Concepts 11.1 Conspecific attraction and conservation
    Applying the Concepts 11.2 Human aggression, testosterone, and sports
    Applying the Concepts 11.3 Reducing duration and intensity of piglet fights
    Toolbox 11.1 The hawk-dove model

    Quantitative Reasoning 11.1 Trout territoriality

    Chapter 12. Mating Behavior
    12.1 Sexual selection favors characteristics that enhance reproductive success
    Why two sexes?
    Bateman's hypothesis and parental investment
    Weapon size and mating success in dung beetles
    Ornaments and mate choice in peafowl
    Male mate choice in pipefish
    The sensory bias hypothesis in guppies
    12.2 Females select males to obtain direct material benefits
    Female choice and nuptial gifts in butterflies
    Female choice and territory quality in lizards
    12.3 Female mate choice can evolve via indirect benefits to offspring
    Fisherian runaway and good genes
    Mate choice for good genes in tree frogs
    Good genes and the Hamilton-Zuk hypothesis
    Mate choice fitness benefits in spiders
    12.4 Sexual selection can also occur after mating
    Mate guarding in warblers
    Sperm competition in tree swallows
    Cryptic female choice
    Inbreeding avoidance via cryptic female choice in spiders
    12.5 Mate choice by females favors alternative reproductive tactics in males
    The evolution of alternative reproductive tactics
    Conditional satellite males in tree frogs
    ESS and sunfish sneaker males
    12.6 Mate choice is affected by the mating decisions of others
    Mate copying in guppies
    Mate copying in fruit flies
    The benefit of mate copying
    Nonindependent mate choice by male mosquitofish
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 12.1 Male mate choice in pipefish
    Scientific Process 12.2 Mate copying in fruit flies
    Applying the Concepts 12.1 Mate choice in conservation breeding programs
    Applying the Concepts 12.2 Human mate choice copying

    Quantitative Reasoning 12.1 Sneaking behavior in New Zealand giraffe weevils

    Chapter 13. Mating Systems
    13.1 Sexual conflict and environmental conditions affect the evolution of mating systems
    The evolution of mating systems
    Mating systems in reed warblers
    13.2 Biparental care favors the evolution of monogamy
    California mouse monogamy
    Monogamy and biparental care in poison frogs
    Monogamy without biparental care in snapping shrimp
    13.3 Polygyny and polyandry evolve when one sex can defend multiple mates or the resources they seek
    Female defense polygyny in horses
    Resource defense polygyny in blackbirds
    Resource defense polygyny in carrion beetles
    Male dominance polygyny: the evolution of leks-hotspots or hotshots?
    Lekking behavior in the great snipe
    Peafowl leks
    Polyandry and sex-role reversal
    13.4 The presence of social associations distinguishes polygynandry from promiscuity
    Polygynandry in European badgers
    Promiscuity and scramble competition in seaweed flies and red squirrels
    13.5 Social and genetic mating systems differ when extra-pair mating occurs
    Extra-pair mating in juncos
    Marmot extra-pair mating
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 13.1 Biparental care and monogamy in poison frogs
    Scientific Process 13.2 Monogamy in snapping shrimp
    Applying the Concepts 13.1 Mating systems and conservation translocation programs
    Applying the Concepts 13.2 Human mating systems
    Toolbox 13.1 DNA fingerprinting

    Quantitative Reasoning 13.1 Mating success of male red-backed fairy-wrens

    Chapter 14. Parental Care
    14.1 Parental care varies among species and reflects life history trade-offs
    Life history variation in fish
    14.2 Sexual conflict is the basis for sex-biased parental care
    Female-biased parental care
    Paternity uncertainty and parental care in boobies
    The evolution of male-only care
    Paternity uncertainty and male-only care in sunfish
    Paternity assurance and male care in water bugs
    14.3 Parental care involves fitness trade-offs between current and future reproduction
    Parent-offspring conflict theory
    Predation risk and parental care in songbirds
    Egg guarding and opportunity costs of parental care in frogs
    Current versus future reproduction in treehoppers
    Incubation of eider eggs as a trade-off
    Brood reduction and parent-offspring conflict
    Hatch asynchrony and brood reduction in blackbirds
    Brood reduction in fur seals
    14.4 Brood parasitism reduces the cost of parental care and can result in a co-evolutionary arms race
    Conspecific brood parasitism in ducks
    Interspecific brood parasitism and co-evolution
    Acceptance or rejection of brown-headed cowbird eggs by hosts
    14.5 Hormones regulate parental care
    Prolactin and maternal care in rats
    Prolactin and incubation in penguins
    Juvenile hormones and parental care in earwigs
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion

    Scientific Process 14.1 Paternity certainty and parental care in bluegill sunfish
    Scientific Process 14.2 Parental care costs in eiders
    Scientific Process 14.3 Brood reduction in blackbirds
    Applying the Concepts 14.1 Human life history trade-off
    Applying the Concepts 14.2 Smallmouth bass defend their nest from exotic predators
    Applying the Concepts 14.3 Food supplementation reduces brood reduction in endangered eagles

    Quantitative Reasoning 14.1 Prey provisioning rates of American kestrals

    Chapter 15. Sociality
    15.1 Sociality can evolve when the fitness advantages of close associations exceed the costs
    Reduced search times for food
    Foraging benefit: Information about distant food locations
    Antipredator benefit of sociality in birds
    Movement benefits: Efficient aerodynamics and hydrodynamics
    Hydrodynamics in schools of juvenile grey mullet
    Social heterosis in ants
    The costs of sociality
    Group size and food competition in red colobus and red-tailed guenons
    Sociality and disease transmission in guppies
    15.2 Dominance hierarchies reduce the social costs of aggression
    Dominance hierarchies and crayfish
    Stable dominance hierarchies in baboons
    15.3 Ecology and phylogeny influence the evolution of sociality
    Evolution of rodent sociality and habitat use
    Body size, diet, and habitat influence sociality in antelope
    15.4 Hormones regulate social behavior
    Social approach behavior and neuropeptides in goldfish
    Mesotocin and pro-social behavior in finches
    Social behavior in seals
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 15.1 Mesotocin and sociality in zebra finches
    Applying the Concepts 15.1 Group aerodynamic advantages in cyclists
    Applying the Concepts 15.2 Group size of social species in captivity

    Quantitative Reasoning 15.1 Benefits of group foraging

    Chapter 16. Cooperative Behavior
    16.1 Inclusive fitness theory explains the evolution of cooperation among related individuals
    Hamilton's rule
    Belding's ground squirrel alarm calls
    Altruism in turkeys
    16.2 Individuals can discriminate kin from non-kin
    Kin discrimination
    Direct familiarization and kin discrimination in sticklebacks
    Indirect familiarization and kin discrimination in cockroaches
    16.3 Cooperative behavior among unrelated individuals involves byproduct mutualisms or reciprocity
    Direct reciprocity
    The prisoner's dilemma
    Tit-for-tat strategy
    Food sharing in vampire bats
    Allogrooming in Japanese macaques
    Tit-for-tat in red-winged blackbirds
    The snowdrift game
    Migrating bald ibis and the snowdrift game
    Indirect reciprocity
    Reputations and cleaner fish
    Reputation formation in great apes
    16.4 Kinship and ecological constraints favor cooperative reproduction
    The evolution of cooperative breeding in vertebrates
    Cooperative breeding in meerkats
    Cooperative reproduction in long-tailed tits
    Helping behavior in Seychelles warblers
    Social queuing in clownfish
    Invertebrate castes
    The evolution of sterile castes
    Haplodiploidy hypothesis
    Kin selection and ecological constraint hypothesis
    Eusociality in sweat bees
    Chapter Summary and Beyond
    Chapter Review
    Critical Thinking and Discussion
    Scientific Process 16.1 Reputation formation in great apes
    Applying the Concepts 16.1 Human altruism and reputations

    Quantitative Reasoning 16.1 Food sharing in killer whales

    Answers to Selected Critical Thinking and Discussion Questions
    Answers to Scientific Process Box "Evaluate" Questions

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