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Index
Half title
Title page
Imprints page
Epigraph
Contents
Contributors
Foreword
Acknowledgments
Part I Overview and behaviors preceding and following initiation of escape
1 Escape behavior: importance, scope, and variables
1.1 Escape, fitness, and predator–prey encounters
1.2 Focus and goals of this book
1.3 Escape theory and data
1.4 The rest: related behaviors, locomotor performance, physiology, genetic and maternal influences, personality difference, best practices for field studies, and conclusions
1.5 A standardized terminology for escape and time spent hiding in refuge
1.5.1 Current ambiguity in escape terminology
1.5.2 Terminology for distance variables and proportions
1.5.3 Terminology for temporal variables
1.5.4 Terminology for directional variables
1.6 Electronic supplementary material
Part II Escape and refuge use: theory and findings for major taxonomic groups
IIa Escape theory
2 Theory: models of escape behavior and refuge use
2.1 Introduction
2.2 First economic models of escape and time spent hiding in refuge
2.2.1 Escape
2.2.2 Time spent hiding in refuge
2.2.3 Assumptions and restrictions
2.3 Optimality models of escape and refuge use
2.4 Comparison of the graphical and optimality models
2.5 Flushing early: effects of starting distance and alert distance on flight initiation distance
2.5.1 Starting distance: an unexpected challenge to economic escape theory
2.5.2 Model of effects of starting distance on flight initiation distance: monitoring costs and spontaneous movements
2.5.3 Starting distance, alert distance, and economic escape
2.5.3.1 Effects of monitoring predators on escape decisions
2.5.3.1.1 Cost of not fleeing
2.5.3.1.2 Cost of fleeing
2.5.3.2 Spontaneous movement
2.5.3.3 Rapid advances in understanding effects of starting distance
2.6 Other approaches to modeling escape decisions and refuge use
2.6.1 Effect of direction of approach by predator on flight decisions for escape to a fixed refuge
2.6.2 Game-theoretical model of escape decisions by cryptic prey
2.6.3 Game-theoretical approach to hiding time in refuge
2.6.4 Stochastic dynamic modeling of fitness consequences of hiding time in refuge
2.7 Behaviors prior to fleeing and flight initiation distance: vigilance, alarm calling, and pursuit-deterrent signaling
2.8 Escape latency
2.9 New model of nearest approach distance by a prey approaching an immobile predator
2.10 Relative movement of prey and predator: escape variables and models
2.11 Conclusions
IIb Escape decisions prior to pursuit
3 Mammals
3.1 Introduction
3.2 Predators of mammals and options for escape
3.3 Scanning for predators, risk assessment, and flight initiation distance
3.4 Factors influencing escape decisions in mammals
3.4.1 Effects of the environment and distance to refuge
3.4.2 Habituation to human disturbances and hunting
3.4.3 Type of disturbance
3.4.4 Effects of group size
3.4.5 Effects of insularity and loss of predators
3.4.6 Social and reproductive effects
3.4.7 Domestication effects
3.5 Case study: reindeer and caribou
3.5.1 Introduction
3.5.2 Habituation in relation to genetics, hunting and tourist activities
3.5.3 Rangifer and predators
3.6 Conservation and management implications
3.7 Foci for the future study of mammal escape behavior
3.8 Conclusions
4 Birds
4.1 Introduction
4.2 Frequency distributions of FID
4.3 FID and components of flight
4.4 Sources of variation in FID
4.4.1 Within- and among-individual variation
4.4.2 Heritability, selection, and response to selection
4.5 Biological causes of variation
4.5.1 Body size
4.5.2 Life history
4.5.3 Urbanization
4.5.4 Song
4.5.5 Hormones and FID
4.5.6 Hunting and disturbance
4.5.7 Diet
4.5.8 Sociality
4.5.9 Predation
4.5.10 Parasitism
4.5.11 Habitat
4.5.12 Range and population size
4.5.13 Dispersal
4.5.14 Overall assessment of effect sizes for different categories of studies
4.6 Assumptions
4.6.1 FID reflects predation risk
4.6.2 Morphological adaptations to FID
4.6.3 Sense organs
4.6.4 Brains and cognition
4.7 Population consequences
4.8 Future prospects
5 Reptiles
5.1 Introduction
5.2 Lizards
5.2.1 Predation risk factors that affect escape during approach
5.2.1.1 Position, habitat, and environmental factors
5.2.1.1.1 Distance to refuge, perch height, and direction to refuge
5.2.1.1.2 Microhabitat
5.2.1.1.3 Habitat openness and exposure
5.2.1.1.4 Temperature
5.2.1.1.5 Wind speed
5.2.1.1.6 Time of day and season
5.2.1.2 Prey traits
5.2.1.2.1 Prey speed, morphology, and body condition
5.2.1.2.2 Habituation
5.2.1.2.3 Prey sex, age, and body size
5.2.1.2.4 Autotomy and tail condition
5.2.1.2.5 Female reproductive condition
5.2.1.2.6 Conspicuousness
5.2.1.2.7 Pursuit-deterrent signaling
5.2.1.3 Predator and approach factors
5.2.1.3.1 Predator approach speed
5.2.1.3.2 Directness of approach
5.2.1.3.3 Predator turn direction
5.2.1.3.4 Approach elevation and sidedness
5.2.1.3.5 Repeated approach and previous captures
5.2.1.3.6 Starting distance
5.2.1.3.7 Sudden shadowing
5.2.1.3.8 Predation pressure
5.2.1.3.9 Predator size and number
5.2.1.3.10 Predator facial exposure, direction of gaze, and eye size
5.2.2 Cost of fleeing factors that affect FID, DF, and related variables
5.2.2.1 Foraging
5.2.2.2 Social costs of fleeing
5.2.3 Importance of costs of not fleeing and costs of fleeing: relative effect sizes
5.2.4 Escape strategy
5.2.5 Latency to flee
5.3 Snakes
5.3.1 FID, distance fled, and probability of fleeing
5.3.2 Refuge selection and entry
5.3.3 Escape strategy
5.4 Crocodilians, sphenodontidans, and turtles
5.5 Future directions
6 Fish and amphibians
6.1 Introduction
6.2 The methodological challenges of working with fish and amphibians
6.2.1 How fish and amphibians perceive the world
6.2.2 Neural control of escape behavior
6.2.3 Logistical issues
6.3 Predictions under the Economic Escape Model: 1. Increased risk of capture
6.3.1 Prey attributes
6.3.1.1 When prey is slower
6.3.1.2 When prey is bright and colorful
6.3.1.3 When prey is distracted
6.3.1.4 When prey has previous experience
6.3.2 When the predator represents more risk
6.3.3 When the environment interferes with prey senses or responses
6.3.3.1 Turbidity (decreased visibility)
6.3.3.2 Greater distance to cover
6.4 Predictions under the Economic Escape Model: 2. increased cost of fleeing
6.4.1 Loss of crypsis
6.4.2 Body size
6.4.3 Opportunity costs
6.5 Predictions under the Economic Escape Model: 3. Effectiveness of alternative defense tactics
6.5.1 Crypsis
6.5.2 Alternative defenses: poison
6.5.3 Armor: spines and plates
6.5.4 Retreat to safer sites
6.6 Predictions under the Economic Escape Model: 4. Group size
6.7 Conclusions
7 Invertebrates
7.1 Introduction
7.2 Measures of invertebrate escape response
7.2.1 Watching and waiting
7.2.1.1 Cessation of movement
7.2.1.2 Falling silent
7.2.1.3 “Positioning†behavior
7.2.2 Fleeing
7.2.2.1 Dropping
7.2.2.2 Taking flight: walking, running, flying, or swimming
7.2.2.2.1 Flight initiation distance (FID)
7.2.2.2.2 Distance fled (DF)
7.2.2.2.3 Flight behavior
7.2.2.3 Retreating to cover
7.3 Factors that influence invertebrate escape response
7.3.1 Food handling
7.3.2 Group living
7.3.3 Effects of body size and age on escape behavior
7.3.4 Effects of temperature on escape behavior
7.4 Conclusions and future research
IIc Escape trajectories and strategies during pursuit
8 Prey behaviors during fleeing: escape trajectories, signaling, and sensory defenses
8.1 Introduction
8.2 Directionality of initial escape responses
8.2.1 The meaning of directionality
8.2.2 Individual characteristics and environmental factors affecting directionality
8.2.3 Should responses with a turn toward a predator be considered mistakes?
8.3 Escape trajectories
8.3.1 Methodological issues
8.3.2 Theory and practice: in which direction do animals flee?
8.3.2.1 Case A: a single, optimal trajectory ()
8.3.2.2 Cases B and C: preferred trajectories () and random ETs within a limited angular sector ()
8.3.2.3 Case D: random trajectories spanning 360° relative to the threat ()
8.3.2.4 Case E: ETs toward a refuge ()
8.3.2.5 Case F: intermediate strategy: away from the threat and toward a refuge ()
8.3.2.6 Overall trends
8.3.3 The relative importance of speed and maneuverability in predators chasing prey
8.3.4 Modification of escape trajectories in the presence of conspecifics
8.4 Sensory defenses and signaling linked to fleeing
8.4.1 Signaling vigor to predators
8.4.2 Cues or signals of physical or chemical defenses revealed during fleeing
8.4.3 Signaling to others during fleeing
8.5 Conclusions
IId Refuge use
9 Hiding time in refuge
9.1 Introduction
9.2 The optimal hiding time
9.3 Experimental tests of the predictions of hiding time models
9.3.1 Factors affecting risk of emergence on hiding time
9.3.1.1 Risk due to the predator’s behavior
9.3.1.2 Risk due to the prey’s characteristics
9.3.1.3 “Waiting games†between predator and prey
9.3.2 Costs of refuge use determining hiding time
9.3.2.1 Loss of foraging opportunities
9.3.2.2 Loss of reproductive opportunities
9.3.2.3 Physiological costs
9.3.2.4 Multiple types of predators and conflicting refuge use
9.4 Hiding behavior when in morphological and constructed refuges
9.5 Hiding under simultaneous risks and costs
9.6 Repeated attacks and multiple hiding decisions
9.6.1 Optimal multiple hiding decisions
9.6.2 Risk assessment affecting multiple hiding decisions
9.6.3 Monitoring from the refuge to determine hiding time
9.6.4 Long-term temporal patterns of risk affecting hiding decisions
9.7 Conclusions and future directions
Part III Related behaviors and other factors influencing escape
10 Vigilance, alarm calling, pursuit deterrence, and predation inspection
10.1 Introduction
10.2 Antipredator vigilance
10.2.1 What is vigilance?
10.2.2 Can detection of predators occur when non-vigilant?
10.2.3 The effect of group size on vigilance
10.2.4 Temporal changes in vigilance
10.2.5 Other factors that affect vigilance
10.3 Alarm calling
10.3.1 Alarm calls can convey information about the type of threat
10.3.2 False alarm calls
10.3.3 Alarm calls and antipredator ploys
10.3.4 Alarm calls and mass flight
10.4 Pursuit-deterrence signals
10.4.1 Perception advertisement
10.4.2 Quality advertisement
10.4.3 Evolution of pursuit-deterrent signals
10.5 Predator inspection
10.5.1 Why inspect predators?
10.5.2 Preferences for partners
10.6 Future directions
11 Determinants of lizard escape performance: decision, motivation, ability, and opportunity
11.1 Introduction
11.2 Locomotor performance affects escape decisions
11.2.1 Temperature effects on fight or flight
11.2.2 Impacts of locomotor performance on flight initiation distance
11.3 Motivation: how do animals decide what to do?
11.4 Escape ability
11.4.1 Body size and shape
11.4.2 Limb and muscle morphology
11.4.3 Escape performance and tail autotomy
11.4.4 How muscle physiology is related to escape ability
11.5 Opportunity: escaping in different habitats
11.5.1 Sandy habitats
11.5.2 Arboreal habitats
11.5.3 Saxicolous habitats
11.5.4 Intermittent locomotion and habitat structure
11.6 Conclusions and future directions
12 Sensory systems and escape behavior
12.1 Sensory systems are at the center of predator–prey interactions
12.2 Steps involved in predator–prey interactions from a sensory perspective
12.2.1 Scanning
12.2.2 Detection
12.2.3 Assessing predation risk
12.2.4 Alert and escape
12.3 Implications for predator–prey interactions
13 The physiology of escape
13.1 Introduction
13.2 Defensive behaviors and physiology
13.2.1 Defensive behaviors (see for summary)
13.2.2 Endocrine roles in defense
13.2.3 Anatomy/neurochemistry of flight-relevant defense systems
13.2.4 Systemic drug effects on defensive behaviors, including flight
13.3 Conclusions
14 Maternal and genetic effects on escape: a prospective review
14.1 Introduction
14.2 Empirical estimates of heritability
14.2.1 Case study 1: Flight in scallops
14.2.2 Case study 2: Flight in garter snakes
14.3 Other evidence for genetic basis of escape behaviors
14.3.1 Family effects
14.3.2 Evidence of heritability from artificially selected lines
14.3.3 Evidence for heritability and local adaptation from population comparisons
14.3.4 Species-level comparisons
14.4 Conclusions based on case studies and other evidence of escape heritability
14.5 Maternal effects
14.5.1 Case study: Egg size and alternative patterns of adaptive escape behavior in side-blotched lizards
14.5.2 Endocrine mechanisms of maternal effects on escape behavior
14.5.3 Other maternal provisioning effects on escape behavior
14.5.4 Incubation regimes and maternal care
14.5.5 Maternal environment and condition effects with unknown mechanisms
14.5.6 Conclusions about maternal effects
14.6 General conclusions
15 The personality of escape
15.1 Introduction
15.2 Is escape behavior a personality trait?
15.2.1 Consistency in the escape and hiding responses
15.2.2 Plasticity and habituation of the escape and hiding response
15.2.3 Correlations of escape and hiding behavior with other behavioral traits
15.3 Evolution of personalities and the optimal escape behavior
15.3.1 State-dependent escape and hiding responses
15.3.2 Life history trade-offs when deciding escape responses
15.3.3 Negative frequency-dependent selection on escape responses
15.4 Future directions
Part IV The application and study of escape
16 Best practice for the study of escape behavior
16.1 Behavior of prey as a trial begins
16.2 Number of humans
16.3 Group size
16.4 Starting distance and alert distance
16.5 Flight initiation distance
16.6 Approach speed and angle
16.7 When to stop approaching
16.8 Data to collect
16.9 Statistical analysis of FID data
16.9.1 Use of SD or AD as covariate or study of interactions involving the SD x FID relationship
16.9.2 Should the FID x SD regression be forced through the origin?
16.9.3 Spontaneous movements and the decision to keep observations of immediate flight
16.9.4 Other statistical issues associated with studying FID
16.10 Comparative studies and meta-analyses: a complementary way to study escape
16.11 Conclusions
17 Afterword
17.1 Introduction
17.2 The importance of studying individuals
17.3 The interplay between interspecific studies and intraspecific studies
17.4 Using physiology to define decision-making mechanisms
17.5 The importance of sensory physiology
17.6 Utility, range, and testability of theory
17.7 Application to wildlife conservation and management
17.8 Conclusions
Index
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