Avoiding attack: the evolutionary ecology of crypsis, warning signals, and mimicry / Graeme D. Ruxton, Thomas N. Sherratt, Michael P. Speed

Incluye bibliografía: páginas 210-239 e índice: páginas 240-249

lntroduction Part I: Avoiding detection.. Chapter 1: Background matching.. 1.1 Why crypsis?.. 1.2 Industrial melanism in Biston betularia.. 1.3 Background is a multivariate entity.. 1.4 Combining background matching with other functions.. 1.5 Flicker fusion.. 1.6 Polymorphism of background matching forms.. 1.6.1 A case study: polymorphism in Cepaea.. 1.6.2 Polymorphism through neutral selection.. 1.6.3 Positive selection for polymorphism.. 1.6.4 Definitions related to frequency-dependent predation.. 1.6.5 Search images.. 1.6.6 Control of search rate.. 1.6.7 Comparing search image and search rate mechanisms.. 1.6.8 Neutral selection again.. 1.7 Coping with multiple backgrounds.. 1.8 Masquerade.. 1.9 Conclusion.. Chapter 2: Disruptive colouration.. 2.1 Introduction.. 2.2 Separating disruptive colouration from background matching.. 2.3 Empirical evidence.. 2.4 Conclusion.. Chapter 3: Countershading and counterillurnination.. 3.1 Introduction.. 3.2 Self-shadow concealment and countershading.. 3.3 Direct empirical tests of the advantages of countersl-iading.. 3.4 Indirect evidence.. 3.4.1 The naked mole-rat.. 3.4.2 Countershading in ungulates.. 3.4.3 Countershading in aquatic environments.. 3.4.4 Counterillumination in marii-ie animals.. 3.5 Countershading in aerial, aquatic, and terrestrial systems.. 3.6 Conclusion.. Chapter 4: Transparency and silvering.. 4.1 Transparent objects still reflect and refract.. 4.2 More reasons why perfect transparency need not translate to perfect crypsis.. 4.2.1 Polarization.. 4.2.2 Other wavelengths of light.. 4.2.3 Snell's window.. 4.3 Imperfect transparency can be effective at low light levels.. 4.4 Some parts of an organism cannot be made transparent.. 4.5 The distribution of transparency across habitats.. 4.6 Silvering as a form of crypsis.. 4.7 Conclusion.. Part II: Avoiding attack after detection.. Chapter 5: Secondary defences.. 5.1 The diversity of secondary defences

5.2 Costs and benefits of some behavioural and morphological secondary defences.. 5.2.1 Behavioural defences.. 5.2.2 Morphological and other mechanical defences.. 5.3 Chemical defences.. 5.3.1 Some characteristics of chemical defences.. 5.3.2 Are chemical defences costly?.. 5.4 Costs, benefits, and forms of defence.. 5.5 The evolution of defences.. 5.5.1 Evolutionary pathways.. 5.5.2 Theoretical approaches to the evolution of defences.. 5.5.3 Formal modelling of the evolution of defences.. 5.6 Summary and conclusion.. Chapter 6: Signalling to predators.. 6.1 Introduction.. 6.2 Signalling that an approaching predator has been detected.. 6.3 Signalling that the prey individual is intrinsically difficult to catch.. 6.4 Summary of theoretical work.. 6.5 Empirical evidence from predators.. 6.5.1 Stotting by gazelle.. 6.5.2 Upright stance by hares.. 6.5.3 Push-up displays by lizards.. 6.5.4 Singing by skylarks.. 6.5.5 Predator inspection behaviour by fish.. 6.5.6 Calling by antelope.. 6.5.7 Fin-flicking behaviour by fish.. 6.6 Studies where predator behaviour is not reported.. 6.6.1 Tail-flicking by rails.. 6.6.2 Tail-signalling by lizards.. 6.6.3 Calling by Diana monkeys.. 6.6.4 Snorting in African bovids.. 6.6.5 Tail-flagging by deer.. 6.6.6 Barking by deer.. 6.7 Conclusion.. Chapter 7: The form and function of warning displays.. 7.1 Characteristics of aposematic warning displays.. 7.1.1 Aposematism does not require complete avoidance by predators.. 7.1.2 Conspicuous animals are not necessarily aposematic.. 7.2 Design of aposematic displays I: why conspicuousness?.. 7.2.1 The opportunity costs of crypsis.. 7.2.2 Forms of secondary defence and the need for conspicuous.. components of warning displays.. 7.3 Design of aposematic displays II: the psychological properties of predators.. 7.3.1 Unlearnt wariness.. 7.3.2 Aposematism and predator learning.. 7.3.3 Memorability.. 7.3.4 Recognition.. 7.3.5 Summary

7.4 Co-evolution: which came first, conspicuousness or special psychological responses to conspicuousness?.. 7.5 Conclusion: designing a warning display.. Chapter 8: The initial evolution of warning displays.. 8.1 The initial evolution of aposematism: the problem.. 8.2 Stochastic-deterministic scenarios.. 8.3 Spatial aggregation.. 8.3.1 Experimental simulations of aggregation effects.. 8.4 More complex population and predator models for aposematism.. 8.5 Individual selection models.. 8.6 Evaluations of predator psychology models.. 8.7 Alternatives to the rare conspicuous mutant scenario.. 8.7.1 Sexual selection.. 8.7.2 Defences, optimal conspicuousness and apparency.. 8.7.3 Aposematism originated to advertise 'visible' defences.. 8.7.4 Facultative, density-dependent aposematism.. 8.7.5 Simultaneous evolution of defence and conspicuousness.. 8.8 Phylogeny and evolutionary history.. 8.9 The evolution of aposematism: a trivial question with interesting answers?.. Chapter 9: The evolution and maintenance of Müllerian mimicry.. 9.1 Where Müllerian mimicry fits in.. 9.2 Chapter outline.. 9.3 A brief early history of Müllerian mimicry.. 9.4 Some potential examples of Müllerian mimicry.. 9.4.1 Neotropical Heliconius butterflies.. 9.4.2 European burnet moths.. 9.4.3 Bumble bees.. 9.4.4 Cotton stainer bugs (genus Dysdercus.. 9.4.5 Poison arrow frogs.. 9.5 Experimental evidence for Müllerian mimicry.. 9.5.1 Direct assessments of the benefits of adopting a common warning signal.. 9.5.2 Proportions of unpalatable prey consurned by naive predators in the course of education.. 9.6 Models of Müllerian mimicry.. 9.7 Questions and controversies.. 9.7.1 Which is the model and which is the mimic?.. 9.7.2 How can mimicry evolve through intermediate stages?.. 9.7.3 Why are mimetic species variable in form between areas?.. 9.7.4 How can multiple Müllerian mimicry rings co-exist?

9.7.5 What is the role of predator generalization in Müllerian mimicry?.. 9.7.6 Why are some Müllerian mimics polymorphic?.. 9.7.7 Do Müllerian mutualists only benefit simply from shared predator education?.. 9.8 Overview.. Part III: Deceiving predators.. Chapter 10: The evolution and maintenance of Batesian mimicry.. 10.1 Scope.. 10.2 Taxonornic distribution of Batesian mimicry.. 10.2.1 Examples of Batesian mimicry.. 10.2.2 Comparative evidence for Batesian mimicry.. 10.3 Experimental evidence for Batesian mimicry and its characteristics.. 10.3.1 Predators learn to avoid noxious models and consequently their palatable mimics.. 10.3.2 Palatable prey altered to resemble an unpalatable species sometimes survive better than mock controls.. 10.3.3 Batesian mimics generally require the presence of the model to gain significant protection.. 10.3.4 The relative (and absolute abundances of the model and mimic affects the rate of predation on these species.. 10.3.5 The distastefulness of the model affects the rate of predation on the model and mimic.. 10.3.6 The model can be simply difficult to catch rather than noxious on capture.. 10.3.7 The success of mimicry is dependent on the availability of alternative prey.. 10.3.8 Mimics do not always have to be perfect replicas to gain protection, particularly when the model is relatively common or highly noxious.. 10.3.9 Frequency-dependent selection on Batesian mimics can lead to mimetic polymorphism.. 10.4 The theory of Batesian mimicry.. 10.5 Questions and controversies.. 10.5.1 Why are not al1 palatable prey Batesian mimics?.. 10.5.2 1s the spatio-temporal coincidence of the models and mimics necessary?.. 10.5.3 Why is Batesian mimicry often limited to one sex?.. 10.5.4 How is mimicry controlled genetically and how can polymorphic mimicry be maintained?.. 10.5.5 Why are imperfect mimics not improved by natural selection?

10.5.6 How does Batesian mimicry evolve, and why do models simply not evolve away from their mimics?.. 10.5.7 What selective factors influence behavioural mimicry?.. 10.6 Overview.. Chapter 11: The relationship between Batesian and Müllerian mimicry.. 11.1 Context.. 11.2 Evidence of interspecific differences in levels of secondary defence.. 11.3 Why should weakly defended mimics increase the likelihood that more highly defended models are attacked?.. 11.3.1 Predator hunger.. 11.3.2 Differences in predatory abilities: the 'Jack Sprat' effect.. 11 3.3 Psychological models.. 11.4 Observational data on the nature of the relationship between Batesian and Müllerian mimicry.. 11.5 Summary.. Chapter 12: Other forms of adaptive resemblance.. 12.1 Overview.. 12.2 Aggressive mimicry.. 12.3 Pollinator (floral mimicry.. 12.4 Intraspecific sexual mimicry.. 12.5 Automimicry.. 12.5.1 The phenomenon of automimicry.. 12.5.2 The challenge to theoreticians.. 12.5.3 Summary.. Chapter 13: Deflection and startling of predators.. 13.1 Deflection defined.. 13.2 Empirical evidence for deflection.. 13.2.1 Lizard tails.. 13.2.2 Tadpole tails.. 13.2.3 Eyespots on fish.. 13.2.4 False head marking on butterflies.. 13.2.5 Weasel tails.. 13.2.6 Summary of empirical evidence for deflective signals.. 13.3 How can deflective marking evolve if they make prey easier for predators to detect?.. 13.Why do predators allow themselves to be deceived?.. 13.5 Startle signals.. 13.5.1 General considerations.. 13.5.2 Distress calls as startle signals.. 13.5.3 Visual startle signals.. 13.5.4 Sound generation by moths attack.. 13.5.5 Summary of empirical evidence.. 13.5.6 Why would predators be startled?.. 13.6 Tonic immobility.. 13.7 Distraction displays.. 13.8 Summary

Chapter 14: General Conclusions.. Appendices.. A: A summary of mathematical and computer models that deal with Müllerian mimicry.. B: A summary of mathematical and computer models that deal with Batesian mimicry.. References.. Author lndex.. Species lndex.. Subject lndex

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 their unprofitability, 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 summarizes some of the latest research into these fascinating adaptations, developing mathematical models where appropriate and making recommendations for the most urgently needed outstanding areas of enquiry. eng