Animal Behavior

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