Animal Behavior Concepts, Methods, and Applications

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.

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.

Preface

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
Behaviorism
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
Features
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
Features
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
Features
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
Features
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
Electroreception
Sharks detect electric fields
Magnetoreception
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
Features
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
Features
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
Teaching
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
Features
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
Features
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
Features
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
Features
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
Features
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
Features
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
Features
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
Features

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
Features
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
Features
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

Glossary
Bibliography
Answers to Selected Critical Thinking and Discussion Questions
Answers to Scientific Process Box "Evaluate" Questions
Credits
Index