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Antipredator Defenses in Birds and Mammals

Illustrated by Sheila Girling
In nature, the ability to defend against predators is fundamental to an animal’s survival. From the giraffes that rely on their spotted coats to blend into the patchy light of their woodland habitats to the South American sea lions that pile themselves in heaps to ward off the killer whales that prey on them in the shallow surf, defense strategies in the animal kingdom are seemingly innumerable.

In Antipredator Defenses in Birds and Mammals, Tim Caro ambitiously synthesizes predator defenses in birds and mammals and integrates all functional and evolutionary perspectives on antipredator defenses that have developed over the last century. Structured chronologically along a hypothetical sequence of predation—Caro evokes a gazelle fawn desperate to survive a cheetah attack to illustrate the continuum of the evolution of antipredator defenses—Antipredator Defenses in Birds and Mammals considers the defenses that prey use to avoid detection by predators; the benefits of living in groups; morphological and behavioral defenses in individuals and groups; and, finally, flight and adaptations of last resort.

Antipredator Defenses in Birds and Mammals will be of interest to both specialists and general readers interested in ecological issues.

592 pages | 15 halftones, 130 line drawings, 90 tables | 6 x 9 | © 2005

Interspecific Interactions

Biological Sciences: Behavioral Biology, Biology--Systematics, Ecology, Evolutionary Biology, Natural History, Physiology, Biomechanics, and Morphology


“This is a field that Caro has devoted much of his career to, and his comfort with the literature shows through. . . .Antipredator Defenses is a major contribution to the field and will be consulted for a long time to come. . . .Caro’s book is a comprehensive survey of the field, with well articulated arguments and balanced views. I recommend it unreservedly.”

Thomas N. Sherratt, | Trends in Ecology and Evolution

"Antipredator Defenses in Birds and Mammals is a comprehensive review of what we know empirically of how animals avoid predation.... [It is] full of fascinating natural history stories that make it an easy and enjoyable read."

Will Cresswell | Ibis

"A reviewer describes the book as ’a wonderful encyclopedic treatment of defenses against predators in mammals and birds,’ but this description is incorrect insofar as it fails to mention that the book is also a great synthesis and magnificently organized. . . . A valuable information bank for any zoologist."

Simone Teelen | Journal of Mammal Evolution

"In-depth reviews of nearly every aspect of prey defenses make this essential reading for researchers and students embarking on studies of prey or predator behavior. The writing is clear and any reader is bound to encounter novel and intriguing material along the way. . . . An important contribution that will help integrate and bolster theoretically and empirically driven work in this field."

Susan Lingle | Ecology

"The work is technically clean and is graced with lovely and topical sketches at the head of each chapter. The treatment is quite nonmathematical and where equations are presented, they are explained quite clearly. . . . This is an important book. . . . Excellent for graduate-level seminars."


"This book is clearly and intelligently written. . . . It could be read with profit by active researchers in predation . . . by those with a more general interest in mammals and birds who are looking for a readable, reliable, and comprehensive reference in tihs area; and by undergraduates in courses related to ecology and animal behavior. . . . I can think of no better recommenation than the considerable number of my colleagues who have bought their own copy after having found themselves reluctant to ask me yet again to lend them mine!"

Graeme D. Ruxton | Bioscience

"An encyclopedic survey that is sure to be a much consulted classic."

Northeastern Naturalist

Table of Contents

Preface, scope, and acknowledgments 

1 Definitions and predator recognition
1.1 Introduction
1.2 The predatory sequence
1.3 Definitions
1.3.a Adaptation and evolution
1.3.b Antipredator terminology
1.4 Ability of prey to recognize predators
1.5 Recognition by young animals
1.5.a Innate recognition
1.5.b Learning to recognize predators
1.6 Relaxed selection
1.7 Observer bias
1.8 Summary 

2 Morphological traits to avoid detection
2.1 Introduction
2.2 Background matching
2.2.a Color resemblance in mammals
2.2.b Color resemblance in birds
2.2.c Color resemblance in birds’ eggs
2.2.d Special resemblance in birds’ nests
2.2.e Melanism
2.2.f Changes in coloration with changing environments
2.2.g Masquerade
2.3 Concealing shadow
2.4 Disruptive coloration
2.5 Apostatic selection
2.6 Summary 

3 Behavioral mechanisms to avoid detection
3.1 Introduction
3.2 Nest site selection in birds
3.2.a Habitat type
3.2.b Distance from edges
3.2.c Habitat patch size
3.2.d Vegetation around the nest site
3.2.e Nest height
3.2.f Proximity to nests
3.2.g Distribution of nests
3.2.h Proximity to social insects
3.3 Behavior reducing the probability of predators detecting nests
3.4 Refuges
3.4.a Physical structures
3.4.b Habitat shifts in rodents
3.4.c Habitat shifts in ungulates
3.5 Reduced activity
3.5.a Hiding in ungulates
3.6 Changes in foraging under risk of predation
3.6.a When to eat
3.6.b Where to eat
3.6.c What to eat
3.6.d How much to eat
3.6.e Effects of age and reproductive condition on risk-sensitive foraging
3.7 Changes in reproduction under risk of predation
3.8 Summary 

4 Vigilance and group size
4.1 Introduction
4.2 Measures of vigilance
4.3 Benefits of individual vigilance
4.4 Costs of individual vigilance
4.5 Effects of group size on vigilance
4.5.a Increased probability of predator detection
4.5.b Reduced individual vigilance
4.5.c Increased foraging
4.6 Why don’t individuals cheat?
4.6.a Predator detection is not collective
4.6.b Vigilant nondetectors are at an advantage
4.6.c Predators select low-vigilance individuals
4.6.d Individuals maintain vigilance so as not to lose group members
4.6.e Multiple attacks are possible
4.7 Vigilance in mixed-species groups
4.8 Summary 

5 Factors affecting vigilance
5.1 Introduction
5.2 Distance from conspecifics and perceived group size
5.3 Position in the group
5.4 Sentinels
5.5 The influence of cover
5.6 Age and parity
5.7 Sex differences and dominance
5.8 Miscellaneous factors
5.9 Predator abundance
5.10 Interspecific differences in vigilance
5.11 Summary 

6 Conspecific warning signals
6.1 Introduction
6.2 Acoustic constraints on alarm calls
6.2.a Localizability
6.2.b Detectability
6.3 Costs of warning signals
6.4 Benefits of warning signals
6.4.a Apparently selfish alarm calls
6.4.b Mutually beneficial alarm calls
6.4.c Altruistic and kin-selected alarm calls
6.5 Alarm calls between species
6.6 Variation in alarm calls
6.6.a Sciurids
6.6.b Birds
6.6.c Primates
6.7 Development of conspecific warning signals
6.7.a Ontogeny of response
6.7.b Ontogeny of alarm calls
6.8 Use of warning signals in deception
6.9 Summary 

7 Signals of unprofitability
7.1 Introduction
7.2 The evolution of aposematism
7.2.a Individual selection
7.2.b Kin selection
7.2.c Synergistic selection
7.3 Mechanisms by which predators select prey
7.3.a Single prey
7.3.b Aggregated prey
7.4 Aposematism in birds
7.4.a Mimicry in birds
7.5 Aposematism in mammals
7.6 Pursuit deterrence
7.6.a Low-cost perception advertisement signals
7.6.b Auditory signals of perception advertisement
7.6.c Inspection as perception advertisement
7.6.d Foot drumming as advertising predator monitoring
7.6.e Stotting as perception and quality advertisement
7.6.f Leaping as quality advertisement
7.6.g Song as quality advertisement
7.6.h Quality advertisement in poikilotherms
7.7 Summary 

8 Antipredator benefits of grouping
8.1 Introduction
8.1.a Definition of groups
8.2 The dilution effect
8.2.a Rates of encounter
8.2.b Reduced risk of capture
8.3 The Trafalgar effect
8.4 The confusion effect
8.4.a Oddity and confusion
8.5 Predator "swamping"
8.5.a Reproductive synchrony
8.6 Miscellaneous mechanisms
8.7 Position in the group
8.7.a Colonially nesting birds
8.7.b Flocks and herds
8.8 Primate groups
8.9 Ecocorrelates of antipredator grouping in homeotherms
8.10 Summary 

9 Morphological and physiological defenses
9.1 Introduction
9.2 Body size
9.2.a Body size and locomotor performance
9.3 Forms of locomotion
9.4 Spines and quills
9.5 Dermal plates and thickened skin
9.6 Weapons used for feeding
9.7 Sexually selected weaponry
9.8 Malodor and unpalatability
9.9 Venom resistance
9.10 Life history characteristics
9.11 Summary 

10 Nest defense
10.1 Introduction
10.2 Scope of nest defense activities
10.2.a The study of nest defense
10.3 Distraction displays
10.4 Costs of nest defense
10.5 Benefits of nest defense
10.5.a Driving predators away
10.5.b Silencing offspring
10.6 Effects of predation risk on nest defense
10.7 Parent’s renesting potential
10.7.a Renesting potential within breeding seasons
10.7.b Renesting potential over lifetimes
10.8 Parental sex
10.9 Parental interactions
10.10 Offspring age
10.10.a Past and future parental investment
10.10.b Changes in offspring vulnerability
10.10.c Revisitation hypothesis
10.11 Offspring number
10.12 Offspring condition
10.12.a Harm-to-offspring hypothesis
10.13 Parental defense in mammals
10.14 Summary 

11 Mobbing and group defense
11.1 Introduction
11.2 Definition of mobbing
11.3 Variation in mobbing behavior
11.4 Costs of mobbing
11.5 Benefits of mobbing
11.5.a Direct benefits: lethal counterattack
11.5.b Direct benefits: the move-on hypothesis
11.5.c Direct benefits: perception advertisement
11.5.d Direct benefits: selfish-herd effect and confusion effect
11.5.e Direct benefits: attract the mightier
11.5.f Indirect benefits: alerting others
11.5.g Indirect benefits: silencing offspring
11.5.h Benefits unclear: cultural transmission
11.5.i Other hypotheses
11.6 Mobbing and group size
11.7 Mobbing and mixed-species associations in birds
11.8 Group defense in mammals
11.8.a Snake-directed behavior in sciurids
11.8.b Protective behavior in ungulates
11.8.c Group attacks in primates
11.9 Summary

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