Avoiding Attack The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry

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Format: Hardcover
Pub. Date: 2005-02-03
Publisher(s): Oxford University Press
List Price: $194.01

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Summary

This book discusses the diversity of mechanisms by which prey avoid attack by predators and questions how such defensive mechanisms have evolved through natural selection. It considers how potential prey avoid detection, how they make themselves unprofitable to attack, how they signal theirunprofitability, and how other species have exploited these signals. Using carefully selected examples drawn from a wide range of species and ecosystems, the authors present a critical analysis of the most important published works in the field. Illustrative examples of camouflage, mimicry and warning signals regularly appear in undergraduate ecology textbooks, but these subjects are rarely considered in depth. This book summarises some of the latest research into these fascinating adaptations, developing mathematical models whereappropriate and making recommendations for the most urgently needed outstanding areas of enquiry.

Author Biography


Graeme Ruxton has co-written two books, both published by Oxford University Press - 'Living in Groups' (2002) for the Oxford Series in Ecology and Evolution, and the textbook 'Elementary Experimental Design for the Life Sciences' (2003). He is also the author of over 100 scientific articles. His background in physics provides particular strength in the functional aspects of signalling systems discussed in this book. Tom Sherratt is the author of nearly 50 scientific papers on subjects ranging from the evolution of co-operation, to the maintenance of imperfect mimicry and the evolution of warning signals. His practical background in both tropical and temperate entomology (principally damselflies and mosquitoes) has been of great value in evaluating empirical work in this broad field, whilst his wide interests in evolutionary biology and foraging theory complement those of his co-authors in placing empirical findings within an appropriate theoretical context. Mike Speed has worked for over a decade on the role of predator behaviour in the generation of insect warning signals. He is consulting editor for the journal Animal Behaviour and a member of the education committee of the Association for the Study of Animal Behaviour. His publications span theoretical and empirical studies of mimicry and aposematism.

Table of Contents

Introduction 1(4)
Part I: Avoiding detection
5(44)
Background matching
7(19)
Why crypsis?
7(2)
Industrial melanism in Biston betularia
9(1)
Background is a multivariate entity
10(1)
Combining background matching with other functions
11(1)
Flicker fusion
12(1)
Polymorphism of background matching forms
12(8)
A case study: polymorphism in Cepaea
13(1)
Polymorphism through neutral selection
13(1)
Positive selection for polymorphism
14(1)
Definitions related to frequency-dependent predation
14(3)
Search images
17(1)
Control of search rate
18(1)
Comparing search image and search rate mechanisms
18(1)
Neutral selection again
19(1)
Coping with multiple backgrounds
20(3)
Masquerade
23(2)
Conclusion
25(1)
Disruptive colouration
26(4)
Introduction
26(1)
Separating disruptive colouration from background matching
27(1)
Empirical evidence
27(2)
Conclusion
29(1)
Countershading and counterillumination
30(8)
Introduction
30(1)
Self-shadow concealment and countershading
30(1)
Direct empirical tests of the advantages of countershading
31(2)
Indirect evidence
33(3)
The naked mole-rat
33(1)
Countershading in ungulates
34(1)
Countershading in aquatic environments
34(1)
Counterillumination in marine animals
35(1)
Countershading in aerial, aquatic, and terrestrial systems
36(1)
Conclusion
37(1)
Transparency and silvering
38(11)
Transparent objects still reflect and refract
38(1)
More reasons why perfect transparency need not translate to perfect crypsis
39(3)
Polarization
39(1)
Other wavelengths of light
40(1)
Snell's window
41(1)
Imperfect transparency can be effective at low light levels
42(1)
Some parts of an organism cannot be made transparent
43(1)
The distribution of transparency across habitats
44(1)
Silvering as a form of crypsis
45(3)
Conclusion
48(1)
Part II: Avoiding attack after detection
49(88)
Secondary defences
51(19)
The diversity of secondary defences
51(2)
Costs and benefits of some behavioural and morphological secondary defences
53(2)
Behavioural defences
53(1)
Morphological and other mechanical defences
54(1)
Chemical defences
55(8)
Some characteristics of chemical defences
56(3)
Are chemical defences costly?
59(4)
Costs, benefits, and forms of defence
63(1)
The evolution of defences
64(4)
Evolutionary pathways
64(1)
Theoretical approaches to the evolution of defences
64(3)
Formal modelling of the evolution of defences
67(1)
Summary and conclusion
68(2)
Signalling to predators
70(12)
Introduction
70(1)
Signalling that an approaching predator has been detected
70(3)
Signalling that the prey individual is intrinsically difficult to catch
73(2)
Summary of theoretical work
75(1)
Empirical evidence from predators
75(4)
Stotting by gazelle
75(1)
Upright stance by hares
76(1)
Push-up displays by lizards
77(1)
Singing by skylarks
77(1)
Predator inspection behaviour by fish
77(1)
Calling by antelope
78(1)
Fin-flicking behaviour by fish
78(1)
Studies where predator behaviour is not reported
79(2)
Tail-flicking by rails
79(1)
Tail-signalling by lizards
80(1)
Calling by Diana monkeys
80(1)
Snorting in African bovids
81(1)
Tail-flagging by deer
81(1)
Barking by deer
81(1)
Conclusion
81(1)
The form and function of warning displays
82(22)
Characteristics of aposematic warning displays
82(3)
Aposematism does not require complete avoidance by predators
84(1)
Conspicuous animals are not necessarily aposematic
84(1)
Design of aposematic displays I: why conspicuousness?
85(4)
The opportunity costs of crypsis
87(1)
Forms of secondary defence and the need for conspicuous components of warning displays
87(2)
Design of aposematic displays II: the psychological properties of predators
89(11)
Unlearnt wariness
90(4)
Aposematism and predator learning
94(3)
Memorability
97(2)
Recognition
99(1)
Summary
100(1)
Co-evolution: which came first, conspicuousness or special psychological responses to conspicuousness?
100(1)
Conclusion: designing a warning display
101(3)
The initial evolution of warning displays
104(11)
The initial evolution of aposematism: the problem
104(1)
Stochastic--deterministic scenarios
105(1)
Spatial aggregation
106(2)
Experimental simulations of aggregation effects
108(1)
More complex population and predator models for aposematism
108(1)
Individual selection models
109(1)
Evaluations of predator psychology models
110(1)
Alternatives to the rare conspicuous mutant scenario
111(2)
Sexual selection
111(1)
Defences, optimal conspicuousness and apparency
112(1)
Aposematism originated to advertise `visible' defences
112(1)
Facultative, density-dependent aposematism
112(1)
Simultaneous evolution of defence and conspicuousness
113(1)
Phylogeny and evolutionary history
113(1)
The evolution of aposematism: a trivial question with interesting answers?
114(1)
The evolution and maintenance of Mullerian mimicry
115(22)
Where Mullerian mimicry fits in
115(1)
Chapter outline
115(1)
A brief early history of Mullerian mimicry
116(2)
Some potential examples of Mullerian mimicry
118(4)
Neotropical Heliconius butterflies
119(1)
European burnet moths
120(1)
Bumble bees
120(2)
Cotton stainer bugs (genus Dysdercus)
122(1)
Poison arrow frogs
122(1)
Experimental evidence for Mullerian mimicry
122(4)
Direct assessments of the benefits of adopting a common warning signal
122(2)
Proportions of unpalatable prey consumed by naive predators in the course of education
124(2)
Models of Mullerian mimicry
126(1)
Questions and controversies
126(10)
Which is the model and which is the mimic?
126(1)
How can mimicry evolve through intermediate stages?
127(2)
Why are mimetic species variable in form between areas?
129(2)
How can multiple Mullerian mimicry rings co-exist?
131(3)
What is the role of predator generalization in Mullerian mimicry?
134(1)
Why are some Mullerian mimics polymorphic?
134(1)
Do Mullerian mutualists only benefit simply from shared predator education?
135(1)
Overview
136(1)
Part III: Deceiving predators
137(65)
The evolution and maintenance of Batesian mimicry
139(25)
Scope
139(1)
Taxonomic distribution of Batesian mimicry
140(2)
Examples of Batesian mimicry
140(1)
Comparative evidence for Batesian mimicry
141(1)
Experimental evidence for Batesian mimicry and its characteristics
142(10)
Predators learn to avoid noxious models and consequently their palatable mimics
142(1)
Palatable prey altered to resemble an unpalatable species sometimes survive better than mock controls
143(1)
Batesian mimics generally require the presence of the model to gain significant protection
144(3)
The relative (and absolute) abundances of the model and mimic affects the rate of predation on these species
147(1)
The distastefulness of the model affects the rate of predation on the model and mimic
148(1)
The model can be simply difficult to catch rather than noxious on capture
148(2)
The success of mimicry is dependent on the availability of alternative prey
150(1)
Mimics do not always have to be perfect replicas to gain protection, particularly when the model is relatively common or highly noxious
150(1)
Frequency-dependent selection on Batesian mimics can lead to mimetic polymorphism
151(1)
The theory of Batesian mimicry
152(2)
Questions and controversies
154(9)
Why are not all palatable prey Batesian mimics?
154(1)
Is the spatio-temporal coincidence of the models and mimics necessary?
155(1)
Why is Batesian mimicry often limited to one sex?
156(2)
How is mimicry controlled genetically and how can polymorphic mimicry be maintained?
158(1)
Why are imperfect mimics not improved by natural selection?
159(2)
How does Batesian mimicry evolve, and why do models simply not evolve away from their mimics?
161(1)
What selective factors influence behavioural mimicry?
162(1)
Overview
163(1)
The relationship between Batesian and Mullerian mimicry
164(8)
Context
164(1)
Evidence of interspecific differences in levels of secondary defence
165(1)
Why should weakly defended mimics increase the likelihood that more highly defended models are attacked?
166(4)
Predator hunger
166(3)
Differences in predatory abilities: the `Jack Sprat' effect
169(1)
Psychological models
169(1)
Observational data on the nature of the relationship between Batesian and Mullerian mimicry
170(1)
Summary
171(1)
Other forms of adaptive resemblance
172(11)
Overview
172(1)
Aggressive mimicry
172(2)
Pollinator (floral) mimicry
174(1)
Intraspecific sexual mimicry
175(1)
Automimicry
176(7)
The phenomenon of automimicry
176(3)
The challenge to theoreticians
179(3)
Summary
182(1)
Deflection and startling of predators
183(17)
Deflection defined
183(1)
Empirical evidence for deflection
183(8)
Lizard tails
183(1)
Tadpole tails
184(1)
Eyespots on fish
185(2)
False head marking on butterflies
187(3)
Weasel tails
190(1)
Summary of empirical evidence for deflective signals
190(1)
How can deflective marking evolve if they make prey easier for predators to detect?
191(1)
Why do predators allow themselves to be deceived?
191(1)
Startle signals
192(5)
General considerations
192(1)
Distress calls as startle signals
193(1)
Visual startle signals
194(1)
Sound generation by moths attacked by bats
195(1)
Summary of empirical evidence
196(1)
Why would predators be startled?
196(1)
Tonic immobility
197(1)
Distraction displays
198(1)
Summary
199(1)
General Conclusions
200(2)
Appendices
202(8)
A: A summary of mathematical and computer models that deal with Mullerian mimicry
B: A summary of mathematical and computer models that deal with Batesian mimicry
References 210(30)
Author Index 240(3)
Species Index 243(5)
Subject Index 248

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