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 hypoth eses 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