B r i e f C o n t e n t s
Chapter 1 The Science of Animal Behavior 2
Chapter 2 Evolution and the Study of Animal Behavior 20
Chapter 3 Methods for Studying Animal Behavior 38
Chapter 4 Behavioral Genetics 56
Chapter 5 Learning and Cognition 78
Chapter 6 Communication 112
Chapter 7 Foraging Behavior 142
Chapter 8 Antipredator Behavior 170
Chapter 9 Dispersal and Migration 196
Chapter 10 Habitat Selection, Territoriality, and Aggression 226
Chapter 11 Mating Behavior 254
Chapter 12 Mating Systems 286
Chapter 13 Parental Care 312
Chapter 14 Social Behavior 338
A p p e n d i x 1 Scientific Literature 372
Appendix 2 Writing a Scientific Paper 374
Appendix 3 Animal Care Information 377 C o n t e n t s
P r e f a c e x x i i i
Chapter 1 The Science of Animal Behavior 2
1.1 Animals and their behavior are an integral part of human society 4 Recognizing and defi ning behavior 5 Measuring behavior: elephant ethograms 5
1.2 Th e scientifi c method is a formalized way of knowing about the natural world 6 Th e importance of hypotheses 7 Th e scientifi c method 7 Correlation and causality 10 Hypotheses and theories 11 Social sciences and the natural sciences 11
1.3 Animal behavior scientists test hypotheses to answer research questions about behavior 12 Hypothesis testing in wolf spiders 12 Negative results and directional hypotheses 13 Generating hypotheses 14 Hypotheses from mathematical models 14
1.4 Anthropomorphic explanations of behavior assign human emotions to animals and can be diffi cult to test 15
1.5 Scientifi c knowledge is generated and communicated to the scientifi c community via peer-reviewed research 17 Th e primary literature 17 Th e secondary literature 18
Chapter Summary and Beyond 19 Q u e s t i o n s 1 9
F e a t u r e s SCIENTIFIC PROCESS 1.1 – Robin abundance and food availability 9 SCIENTIFIC PROCESS 1.2 – Robin abundance and predators 10 TOOLBOX 1.1 – Scientifi c literacy 18 APPLYING THE CONCEPTS 1.1 – What is behind the “guilty look” in dogs? 16 x Contents
C h a p t e r 2 Evolution and the Study of Animal Behavior 20
2.1 Evolution by natural selection favors behavioral adaptations that enhance fi tness 22 M e a s u r e s o f h e r i t a b i l i t y 2 3 Great tit exploratory behavior 24 Variation within a population 25 Fitness and adaptation 26
2.2 Modes of natural selection describe population changes 28 Directional selection in juvenile ornate tree lizards 28 Disruptive selection in spadefoot toad tadpoles 30 Stabilizing selection in juvenile convict cichlids 31 Studying adaptation: the cost-benefi t approach 32
2.3 Individual and group selection have been used to explain cooperation 32
2.4 Sexual selection is a form of natural selection that focuses on the reproductive fi tness of individuals 35 Sexual selection in house fi nches 35
Chapter Summary and Beyond 37 Q u e s t i o n s 3 7
F e a t u r e s SCIENTIFIC PROCESS 2.1 – Heritability of great tit exploratory behavior 25 SCIENTIFIC PROCESS 2.2 – Stabilizing selection on territory size in cichlids 33 TOOLBOX 2.1 – Genetics primer 23 TOOLBOX 2.2 – Frequency-dependent selection 27 TOOLBOX 2.3 – Game theory 34 APPLYING THE CONCEPTS 2.1 – Do lemmings commit suicide? 35
C h a p t e r 3 Methods for Studying Animal Behavior 38
3.1 Scientists study both the proximate mechanisms that generate behavior and the ultimate reasons why the behavior evolved 40
3.2 Researchers use observational, experimental, and comparative methods to study behavior 41 Th e observational method 41 Th e observational method and reproductive energetics of chimpanzees 42 Th e experimental method 43 Th e experimental method and jumping tadpoles 44 Th e comparative method 46 Th e comparative method and the evolution of burrowing behavior in mice 46 Contents xi
3.3 Researchers have examined animal behavior from a variety of perspectives over time 48 Darwin and adaptation 49 Early comparative psychology 49 Comparative psychology in North America 50 B e h a v i o r i s m 5 0 C l a s s i c a l e t h o l o g y 5 1 I n t e r d i s c i p l i n a r y a p p r o a c h e s 5 1
3.4 Animal behavior research requires ethical animal use 52 How research can aff ect animals 52 Sources of ethical standards 53 Th e three Rs 53
Chapter Summary and Beyond 55 Q u e s t i o n s 5 5
F e a t u r e s SCIENTIFIC PROCESS 3.1 – Jumping tadpoles 45 TOOLBOX 3.1 – Animal sampling techniques 44 TOOLBOX 3.2 – Creating phylogenies 47 TOOLBOX 3.3 – Describing and summarizing your data 54 APPLYING THE CONCEPTS 3.1 – Project Seahorse 42
C h a p t e r 4 Behavioral Genetics 56
4.1 Behavioral variation is associated with genetic variation 58 Th e search for a genetic basis of behavior 58 Behavioral diff erences between wild-type and mutant-type fruit fl ies 59 Major and minor genes 60 QTL mapping to identify genes associated with behavior 60 QTL mapping for aphid feeding behavior 60 Fire ant genotype and social organization 61 Experimental manipulation of gene function: knockout studies 63 Anxiety-related behavior and knockout of a hormone receptor in mice 63
4.2 Th e environment infl uences gene expression and behavior 65 H e r i t a b i l i t y 6 5 Environmental eff ects on zebrafi sh aggression 67 Social environment and gene expression in fruit fl ies 67 Social environment and birdsong development 69 Social environment and gene expression in birds 70 Gene-environment interactions 71 Rover and sitt er foraging behavior in fruit fl ies 71 xii Contents
4.3 Genes can limit behavioral fl exibility 74 Bold-shy personalities in streamside salamanders 74 Animal personalities: a model with fi tness trade-off s 76
Chapter Summary and Beyond 77 Q u e s t i o n s 7 7
F e a t u r e s SCIENTIFIC PROCESS 4.1 – Environmental eff ects on zebrafi sh aggression 68 SCIENTIFIC PROCESS 4.2 – Salamander personalities 75 TOOLBOX 4.1 – Molecular techniques 62 TOOLBOX 4.2 – Methods for studying behavior under stress 64 APPLYING THE CONCEPTS 4.1 – Dog behavior heritability 66
C h a p t e r 5 Learning and Cognition 78
5.1 Learning allows animals to adapt to their environment 80 Learning as an adaptation in juncos 80 Evolution of learning 80 Green frog habituation to intruder vocalizations 82
5.2 Learning is associated with neurological changes 83 N e u r o t r a n s m i tt ers and learning in chicks 84 Dendritic spines and learning in mice 85 Avian memory of stored food 87
5.3 Animals learn stimulus-response associations 89 C l a s s i c a l c o n d i t i o n i n g 8 9 Pavlovian conditioning for mating opportunities in Japanese quail 90 Fish learn novel predators 91 O p e r a n t c o n d i t i o n i n g 9 2 Learning curves in macaques 94 Trial-and-error learning in bees 94
5.4 Social interactions facilitate learning 97 Ptarmigan chicks learn their diet 97 Prairie dogs learn about predators 98 Learning the location of food patches 99 Social information use in sticklebacks 100 Public information use in starlings 101
5.5 Social learning can lead to the development of animal traditions and culture 103 Behavioral tradition in wrasse 104 Contents xiii
5.6 Some animals can use mental representations to solve complex problems 105 History of animal cognition research 106 Elephants and insight learning 107 Numerical competency in New Zealand robins 107 Cognition and brain architecture in birds 108
Chapter Summary and Beyond 110 Questions 111
F e a t u r e s SCIENTIFIC PROCESS 5.1 – Brain structure and food hoarding 89 SCIENTIFIC PROCESS 5.2 – Fish learn predators 92 SCIENTIFIC PROCESS 5.3 – Starlings use public information 102 TOOLBOX 5.1 – Neurobiology primer 109 APPLYING THE CONCEPTS 5.1 – Operation migration and imprinting 85 APPLYING THE CONCEPTS 5.2 – Dog training 93 APPLYING THE CONCEPTS 5.3 – Are you smarter than a pigeon? 95
C h a p t e r 6 Communication 112
6.1 Communication occurs when a specialized signal from one individual infl uences the behavior of another 114 Honeybees and the waggle dance 114 Odor or the dance in bees 115 Auditory signals: alarm calls 116 Titmouse alarm calls 117 Information or infl uence? 117
6.2 Signals are perceived by sensory systems and infl uenced by the environment 118 C h e m o r e c e p t i o n 1 1 8 A n t p h e r o m o n e s 1 1 9 P h o t o r e c e p t o r s 1 1 9 Habitat structure and visual signals in birds 120 Auditory receptors 122 Habitat structure and bowerbird vocal signals 123
6.3 Signals can accurately indicate signaler phenotype and environmental conditions 125 Signals as accurate indicators: theory 125 Aposematic coloration in frogs 126 Courtship signaling in spiders 127 Aggressive display and male condition in fi ghting fi sh 129
6.4 Signals can be inaccurate indicators when the fi tness interests of signaler and receiver diff er 129 Batesian mimicry and Ensatina salamanders 131 Aggressive mimicry in fangblenny fi sh 132 Intraspecifi c deception: false alarm calls 133 xiv Contents
Topi antelope false alarm calls 133 Capuchin monkeys and inaccurate signals 135
6.5 Communication networks aff ect signaler and receiver behavior 137 Squirrel eavesdropping 137 Eavesdropping in túngara frogs 138 Audience eff ects in fi ghting fi sh 139
Chapter Summary and Beyond 140 Questions 141
F e a t u r e s SCIENTIFIC PROCESS 6.1 – Chemical signals in ants 120 SCIENTIFIC PROCESS 6.2 – Signaling in male wolf spiders 128 SCIENTIFIC PROCESS 6.3 – Fighting fi sh opercular display 130 APPLYING THE CONCEPTS 6.1 – Pheromones and pest control 119 APPLYING THE CONCEPTS 6.2 – Urban sounds aff ect signal production 122
C h a p t e r 7 Foraging Behavior 142
7.1 Animals fi nd food using a variety of sensory modalities 144 Catfi sh track the wake of their prey 144 Bees use multiple senses to enhance foraging effi ciency 147 Gray mouse lemurs use multiple senses to fi nd food 148
7.2 Visual predators fi nd cryptic prey more eff ectively by learning a search image 149 Cryptic coloration reduces predator effi ciency in trout 150 Blue jays use a search image to fi nd prey 150
7.3 Th e optimal diet model predicts the food types an animal should include in its diet 152 Th e diet model 152 A graphical solution 153 Diet choice in northwestern crows 155 Ant foraging: the eff ect of nutrients 157
7.4 Th e optimal patch-use model predicts how long a forager should exploit a food patch 158 Th e optimal patch-use model 158 Patch use by ruddy ducks 160 Optimal patch model with multiple costs 161 Fruit bats foraging on heterogeneous patches 161 Gerbil foraging with variable predation costs 163 Incomplete information and food patch estimation 164 Bayesian foraging bumblebees 166
Chapter Summary and Beyond 168 Questions 168 Contents xv
F e a t u r e s SCIENTIFIC PROCESS 7.1 – Prey detection by gray mouse lemurs 149 SCIENTIFIC PROCESS 7.2 – Cryptic prey reduces predator effi ciency 151 SCIENTIFIC PROCESS 7.3 – Patch use by fruit bats 162 TOOLBOX 7.1 – Mathematical solution to the optimal diet model 156 APPLYING THE CONCEPTS 7.1 – GUDs and conservation 165
C h a p t e r 8 Antipredator Behavior 170
8.1 Animals modify their behavior to reduce predation risk 172 Predator avoidance by cryptic coloration in crabs 172 Predators and reduced activity in lizards 174 Prey take evasive action when detected 175
8.2 Many behaviors represent adaptive trade-off s involving predation risk 175 Increased vigilance decreases feeding time 176 Vigilance and predation risk in elk 177 Energy intake versus safety in squirrels 178 Rich but risky 180 Environmental conditions and predation risk in foraging redshanks 180 Predation risk and patch quality in ants 182 Mating near predators in water striders 184 Mating and refuge use in fi ddler crabs 185
8.3 Living in groups can reduce predation risk 186 Th e dilution eff ect and killifi sh 186 Th e selfi sh herd and vigilance behavior 187 Group size eff ect and the selfi sh herd hypothesis in doves 188
8.4 Some animals interact with predators to deter att ack 189 Predator harassment in ground squirrels 190 Mobbing owl predators 192 Pursuit deterrence and alarm signal hypotheses 192 Tail-fl agging behavior in deer 193
Chapter Summary and Beyond 194 Questions 195
F e a t u r e s SCIENTIFIC PROCESS 8.1 – Feeding trade-off in redshanks 181 SCIENTIFIC PROCESS 8.2 – Mating behavior trade-off in water striders 185 SCIENTIFIC PROCESS 8.3 – Predator harassment by California ground squirrels 191 APPLYING THE CONCEPTS 8.1 – Mitigating crop damage by manipulating predation risk 176 xvi Contents
C h a p t e r 9 Dispersal and Migration 196
9.1 Dispersal reduces competition and inbreeding 198 Dispersal in adult springtails 198 Natal dispersal in northern goshawks 199 Inbreeding avoidance in voles 200
9.2 Reproductive success aff ects breeding dispersal 202 Breeding dispersal in dragonfl ies 202 Public information from conspecifi cs aff ects breeding dispersal 204 K i tt iwakes and public information 204
9.3 Animals migrate in response to changes in the environment 205 Migration and changing resources 206 Resource variation and migration in neotropical birds 207 Th e evolution of migration 208 Competition and migratory behavior of newts 209 Maintenance of polymorphism in migratory behavior 210 Competition and migratory behavior of dippers 211
9.4 Animals use multiple compass systems to determine direction 212 Th e sun compass in monarch butt erfl ies 213 Antennae and the sun compass system in monarchs 216 Th e magnetic compass in sea turtles 216 M a g n e t o r e c e p t i o n 2 1 8 Multimodal orientation 219
9.5 Bicoordinate navigation allows individuals to identify their location relative to a goal 220 Bicoordinate navigation and magnetic maps in sea turtles 220 Bicoordinate navigation in birds 221
Chapter Summary and Beyond 225 Questions 225
F e a t u r e s SCIENTIFIC PROCESS 9.1 – Breeding dispersal in dragonfl ies 203 SCIENTIFIC PROCESS 9.2 – Th e role of the antennae in the monarch butt erfl y sun compass 217 TOOLBOX 9.1 – Emlen funnels 224 APPLYING THE CONCEPTS 9.1 – Bird migration and global climate change 207 APPLYING THE CONCEPTS 9.2 – Citizen scientists track fall migration fl yways of monarch butt erfl ies 218
C h a p t e r 1 0 Habitat Selection, Territoriality, and Aggression 226
10.1 Resource availability and the presence of others infl uence habitat selection 228 Th e ideal free distribution model 228 Th e ideal free distribution model and guppies 229 Th e ideal free distribution model and pike 231 Cuckoos assess habitat quality 232 Contents xvii
C o n s p e c i fi c a tt raction 233 C o n s p e c i fi c a tt raction and Allee eff ects in grasshoppers 234 Conspecifi c cueing in American redstarts 236
10.2 Individual condition and environmental factors aff ect territoriality 237 Body condition aff ects territoriality in damselfl ies 237 Environmental factors and territory size in kites 239
10.3 Th e decisions of opponents and resource value aff ect fi ghting behavior 240 Th e hawk-dove model 241 Th e sequential assessment model: fi ddler crab contests 241 Resource value and fi ghting behavior in penguins 245 Salamander fi ghts for mates 246
10.4 Hormones infl uence aggression 248 Winner-challenge eff ect in the California mouse 249 Challenge hypothesis and bystanders in fi sh 250
Chapter Summary and Beyond 252 Questions 252
F e a t u r e s SCIENTIFIC PROCESS 10.1 – Ideal free guppies 230 SCIENTIFIC PROCESS 10.2 – Conspecifi c att raction in grasshoppers 235 SCIENTIFIC PROCESS 10.3 – Sequential assessment in crab fi ghts 244 TOOLBOX 10.1 – Th e hawk-dove model 242 APPLYING THE CONCEPTS 10.1 – Conspecifi c att raction and conservation 238 APPLYING THE CONCEPTS 10.2 – Reducing duration and intensity of piglet fi ghts 247 APPLYING THE CONCEPTS 10.3 – Sports, aggression, and testosterone 251
C h a p t e r 1 1 Mating Behavior 254
11.1 Sexual selection favors characteristics that enhance reproductive success 256 Why two sexes? 257 Bateman’s hypothesis and parental investment 257 Antlers as weapons in red deer 258 Weapon size and mating success in dung beetles 259 Ornaments and mate choice in peafowl 260 Th e origin of sexually selected traits: the sensory bias hypothesis in guppies 262 Male mate choice in pipefi sh 263
11.2 Females select males to obtain direct material benefi ts 265 Female choice and nuptial gift s in fi refl ies 265 Female choice and territory quality in lizards 267
11.3 Female mate choice can evolve via indirect benefi ts to off spring 267 Fisherian runaway and good genes 268 Mate choice for good genes in tree frogs 269 xviii Contents
Good genes and immune system function in birds 270 Mate choice fi tness benefi ts in spiders 271
11.4 Sexual selection can also occur aft er mating 273 Mate guarding in warblers 273 Sperm competition in tree swallows 274 Cryptic female choice 275 Inbreeding avoidance via cryptic female choice in spiders 275
11.5 Mate choice by females favors alternative reproductive tactics in males 276 Th e evolution of alternative reproductive tactics 277 Conditional satellite males in tree frogs 277 ESS and sunfi sh sneaker males 278
11.6 Mate choice is aff ected by the mating decisions of others 280 Mate copying in guppies 280 Mate copying in fruit fl ies 281 Th e benefi t of mate copying 281 Nonindependent mate choice by male mosquitofi sh 283
Chapter Summary and Beyond 284 Questions 285
F e a t u r e s SCIENTIFIC PROCESS 11.1 – Male mate choice in pipefi sh 264 SCIENTIFIC PROCESS 11.2 – Mate copying in fruit fl ies 282 APPLYING THE CONCEPTS 11.1 – Mate choice in conservation breeding programs 272
C h a p t e r 1 2 Mating Systems 286
12.1 Sexual confl ict and environmental conditions aff ect the evolution of mating systems 288 Th e evolution of social mating systems 288 Mating systems in reed warblers 291
12.2 Monogamy oft en evolves when biparental care is required to raise off spring 294 California mouse monogamy 294 Monogamy and biparental care in poison frogs 294 Monogamy without biparental care: snapping shrimp 296
12.3 Polygyny and polyandry evolve when one sex can defend multiple mates or the resources they seek 298 Female defense polygyny in horses 298 Resource defense polygyny in blackbirds 299 Resource defense polygyny in carrion beetles 301 Male dominance polygyny: the evolution of leks—hotspots or hotshots? 301 Lekking behavior in the great snipe 303 P e a f o w l l e k s 3 0 3 Polyandry and sex-role reversal 304 Contents xix
12.4 Th e presence of social associations distinguishes polygynandry from promiscuity 305 Polygynandry in European badgers 306 Promiscuity and scramble competition: seaweed fl ies and red squirrels 307
12.5 Social and genetic mating systems diff er when extra-pair mating occurs 308 Extra-pair mating in juncos 309 M a r m o t e x t r a - p a i r m a t i n g 3 1 0
Chapter Summary and Beyond 311 Questions 311
F e a t u r e s SCIENTIFIC PROCESS 12.1 – Biparental care and monogamy in poison frogs 295 TOOLBOX 12.1 – DNA fi ngerprinting 302 APPLYING THE CONCEPTS 12.1 – Mating systems and conservation translocation programs 290
C h a p t e r 1 3 Parental Care 312
13.1 Parental care varies among species and refl ects life history trade-off s 314 Life history variation in fi sh 315
13.2 Sexual confl ict is the basis for sex-biased parental care 316 Female-biased parental care 316 Paternity uncertainty and parental care in boobies 317 Th e evolution of male-only care 317 Paternity uncertainty and male-only care in sunfi sh 318 Paternity assurance and male care in water bugs 320
13.3 Hormones regulate parental care 320 Prolactin and maternal care in rats 321 Prolactin and incubation in penguins 322 Juvenile hormones and parental care in earwigs 323
13.4 Parental care involves fi tness trade-off s between current and future reproduction 324 Predation risk and parental care in golden egg bugs 324 Egg guarding and opportunity costs of parental care in frogs 325 Parent-off spring confl ict theory 326 Parental care trade-off in treehoppers 327 Incubation of eider eggs as a trade-off 329 Brood reduction and parent-off spring confl ict 331 Hatch asynchrony and brood reduction in blackbirds 331 Brood reduction in fur seals 333 Brood parasitism 334 Conspecifi c brood parasitism in ducks 335 xx Contents
Chapter Summary and Beyond 336 Questions 336
F e a t u r e s SCIENTIFIC PROCESS 13.1 – Paternity certainty and parental care in bluegill sunfi sh 319 SCIENTIFIC PROCESS 13.2 – Parental care costs in eiders 329 SCIENTIFIC PROCESS 13.3 – Brood reduction in blackbirds 332 APPLYING THE CONCEPTS 13.1 – Smallmouth bass defend their nest from exotic predators 330 APPLYING THE CONCEPTS 13.2 – Food supplementation reduces brood reduction in endangered eagles 334
C h a p t e r 1 4 Social Behavior 338
14.1 Sociality evolves when the net benefi ts of close associations exceed the costs 340 Foraging benefi ts: reduced search times for food in minnows 341 Foraging benefi ts: increased diet breadth in coyotes 342 Antipredator benefi ts in honeycreepers 343 Aerodynamic benefi t: reduced cost of movement in pelicans 344 Th e costs of sociality 344 Competition in schooling fi sh 345 Group size and competition in primates 345 Sociality and disease transmission in guppies 346
14.2 Dominance hierarchies within groups reduce aggression 348 Dominance hierarchies and crayfi sh 348 Stable dominance hierarchies in baboons 349 Social queuing in dominance hierarchies in clownfi sh 351
14.3 Helping behavior, or altruism, is oft en directed toward close kin 352 Hamilton’s rule 352 Belding ground squirrel alarm calls 353 Altruism and helping at the nest in birds 354 Altruism in turkeys 355
14.4 Some species exhibit extreme altruism in the form of eusociality 357 Eusociality and haplodiploidy 357 Eusociality in ants 358 Evolution of eusociality and kin selection in Hymenoptera 359
14.5 Helping between unrelated individuals requires reciprocity 359 Th e prisoner’s dilemma 361 Reciprocal altruism in vampire bats 362 Allogrooming in Japanese macaques 363 Tit-for-tat in red-winged blackbirds 365 Indirect reciprocity 366 Contents xxi
Reputations and cleaner fi sh 366 Chimpanzee image scores 367
Chapter Summary and Beyond 370 Questions 370
F e a t u r e s SCIENTIFIC PROCESS 14.1 – Chimpanzee image scoring 368 APPLYING THE CONCEPTS 14.1 – Group size of social species in captivity 348 APPLYING THE CONCEPTS 14.2 – Are naked mole rats eusocial? 358 APPLYING THE CONCEPTS 14.3 – Human altruism and reputations 369
A p p e n d i x e s A p p e n d i x 1 — S c i e n t i fi c L i t e r a t u r e 3 72 Appendix 2—Writing a Scientifi c Paper 374 Appendix 3—Animal Care Information 377
G l o s s a r y 3 7 9 B i b l i o g r a p h y 3 9 1 Answers to Selected Questions 419 Credits 423 I n d e x 4 2 7