An Introduction to Behavioural Ecology E-Book

Nicholas B. Davies

• Herausgeber: John Wiley & Sons
• Kategorie: Wissenschaft und neue Technologien
• Sprache: Englisch

Seitenzahl: 1144

Veröffentlichungsjahr: 2012

Beschreibung

This textbook helped to define the field of Behavioural Ecology. In this fourth edition the text has been completely revised, with new chapters and many new illustrations and full colour photographs. The theme, once again, is the influence of natural selection on behaviour – an animal's struggle to survive and reproduce by exploiting and competing for resources, avoiding predators, selecting mates and caring for offspring, – and how animal societies reflect both cooperation and conflict among individuals.

Stuart A. West has joined as a co-author bringing his own perspectives and work on microbial systems into the book.

Written in the same engaging and lucid style as the previous editions, the authors explain the latest theoretical ideas using examples from micro-organisms, invertebrates and vertebrates. There are boxed sections for some topics and marginal notes help guide the reader. The book is essential reading for students of behavioural ecology, animal behaviour and evolutionary biology.

Key Features:

• Long-awaited new edition of a field-defining textbook
• New chapters, illustrations and colour photographs
• New co-author
• Focuses on the influence of natural selection on behavior, and how animal societies reflect both cooperation and conflict among individuals

"The long-awaited update to a classic in this field is now here, presenting new directions in thinking and addressing burning questions. Richly informed by progress in many other disciplines, such as sensory physiology, genetics and evolutionary theory, it marks the emergence of behavioural ecology as a fully fledged discipline.... This is a marvellous book, written in a lucid style. A must-read for those in the field, it is also a cornucopia of new thinking for anyone interested in evolution and behaviour."
—Manfred Milinski, Nature, 2012

Leseprobe

Contents

Preface

Acknowledgements

CHAPTER 1 Natural Selection, Ecology and Behaviour

Watching and wondering

Natural selection

Genes and behaviour

Selfish individuals or group advantage?

Phenotypic plasticity: climate change and breeding times

Behaviour, ecology and evolution

Summary

Further reading

CHAPTER 2 Testing Hypotheses in Behavioural Ecology

The comparative approach

Breeding behaviour of gulls in relation to predation risk

Social organization of weaver birds

Social organization in African ungulates

Limitations of early comparative studies

Comparative approach to primate ecology and behaviour

Using phylogenies in comparative analysis

The comparative approach reviewed

Experimental studies of adaptation

Summary

Further reading

CHAPTER 3 Economic Decisions and the Individual

The economics of carrying a load

The economics of prey choice

Sampling and information

The risk of starvation

Environmental variability, body reserves and food storing

Food storing birds: from behavioural ecology to neuroscience

The evolution of cognition

Feeding and danger: a trade-off

Social learning

Optimality models and behaviour: an overview

Summary

Further reading

CHAPTER 4 Predators versus Prey: Evolutionary Arms Races

Red Queen evolution

Predators versus cryptic prey

Enhancing camouflage

Warning colouration: aposematism

Mimicry

Trade-offs in prey defences

Cuckoos versus hosts

Summary

Further reading

CHAPTER 5 Competing for Resources

The Hawk–Dove game

Competition by exploitation: the ideal free distribution

Competition by resource defence: the despotic distribution

The ideal free distribution with unequal competitors

The economics of resource defence

Producers and scroungers

Alternative mating strategies and tactics

ESS thinking

Animal personalities

Summary

Further reading

CHAPTER 6 Living in Groups

How grouping can reduce predation

How grouping can improve foraging

Evolution of group living: shoaling in guppies

Group size and skew

Group decision making

Summary

Further reading

Chapter 7 Sexual Selection, Sperm Competition and Sexual Conflict

Males and females

Parental investment and sexual competition

Why do females invest more in offspring care than do males?

Evidence for sexual selection

Why are females choosy?

Genetic benefits from female choice: two hypotheses

Testing the hypotheses for genetic benefits

Sexual selection in females and male choice

Sex differences in competition

Sperm competition

Constraints on mate choice and extra-pair matings

Sexual conflict

Sexual conflict: who wins?

Chase-away sexual selection

Summary

Further reading

CHAPTER 8 Parental Care and Family Conflicts

Evolution of parental care

Parental investment: a parent’s optimum

Varying care in relation to costs and benefits

Sexual conflict

Sibling rivalry and parent–offspring conflict: theory

Sibling rivalry: evidence

Parent–offspring conflict: evidence

Brood parasites

Summary

Further reading

CHAPTER 9 Mating Systems

Mating systems with no male parental care

Mating systems with male parental care

A hierarchical approach to mating system diversity

Summary

Further reading

CHAPTER 10 Sex Allocation

Fisher’s theory of equal investment

Sex allocation when relatives interact

Sex allocation in variable environments

Selfish sex ratio distorters

Summary

Further reading

CHAPTER 11 Social Behaviours: Altruism to Spite

Kin selection and inclusive fitness

Hamilton’s rule

How do individuals recognize kin?

Kin selection doesn’t need kin discrimination

Selfish restraint and kin selection

Spite

Summary

Further reading

CHAPTER 12 Cooperation

What is cooperation?

Free riding and the problem of cooperation

Solving the problem of cooperation

Kin selection

Hidden benefits

By-product benefit

Reciprocity

Enforcement

A case study – the Seychelles Warbler

Manipulation

Summary

Further reading

CHAPTER 13 Altruism and Conflict in the Social Insects

The social insects

The life cycle and natural history of a social insect

The economics of eusociality

The pathway to eusociality

The haplodiploidy hypothesis

The monogamy hypothesis

The ecological benefits of cooperation

Conflict within insect societies

Conflict over the sex ratio in the social hymenoptera

Worker policing in the social hymenoptera

Superorganisms

Comparison of vertebrates with insects

Summary

Further reading

CHAPTER 14 Communication and Signals

The types of communication

The problem of signal reliability

Indices

Handicaps

Common interest

Human language

Dishonest signals

Summary

Further reading

CHAPTER 15 Conclusion

How plausible are our main premises?

Causal and functional explanations

A final comment

Summary

Further reading

References

Index

COMPANION WEBSITE

This book is accompanied by a companion website:

    www.wiley.com/go/davies/behaviouralecology

With figures and tables from the book for downloading

This edition first published 2012© 2012 by Nicholas B. Davies, John R. Krebs and Stuart A. West

Previous editions: © 1981, 1987, 1993 by Blackwell Science Ltd.

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Preface

Photo © osf.co.uk.

In this Preface, we summarise the history and organisation of this book. The previous editions, by John Krebs and Nick Davies (1981, 1987, 1993) celebrated the early years of behavioural ecology. Our aim was to understand how behaviour evolves in the natural world. This requires links between studies of behaviour, evolution and ecology. The link with evolution is central because we expect natural selection to favour those behaviour patterns which maximise an individual’s chances of surviving and passing copies of its genes on to future generations. The link with ecology comes in because ecology sets the stage on which individuals play their behaviour, so the best way to behave depends on ecological selection pressures, such as the distribution in space and time of food, enemies and places to live. The social environment will be important too, because individuals will often have to compete for scarce resources. So we need to consider how behaviour evolves when there are social interactions, with the potential for both conflict and cooperation.

This new edition celebrates a maturing and flourishing field, with exciting new links being forged with other disciplines. We now have three co-authors. John was an adviser for Nick’s doctorate research in Oxford and Nick, in turn, lectured to Stu in undergraduate courses in Cambridge. So we span three (short!) academic generations and we have all enjoyed learning from each other during the preparation of this book. All the chapters have been heavily revised or completely reorganised and re-written to incorporate the many new ideas and examples which have emerged since the last edition - in some cases overturning what used to be the conventional wisdom. The central themes remain: a reductionist approach to consider the costs and benefits of decision making and how trade-offs are resolved by selection; a “gene’s eye” view of behaviour; and a game theoretic approach to analyse the resolution of conflicts of interest. John and Nick remember the early days when Bill Hamilton, Robert Trivers and John Maynard Smith began to explore the ideas of kin selection, family conflicts and game theory for the analysis of conflict resolution, and when Richard Dawkins was testing drafts of The Selfish Gene as undergraduate lectures in Oxford. It is exciting to see how these ideas have evolved to stimulate the new research we discuss here. Throughout the book, we emphasise the theoretical background, but we prefer to develop the theory with examples rather than with abstract arguments. Some of the more complicated arguments are presented in boxes.

Chapter 1 begins with “watching and wondering” as we introduce how to frame different kinds of questions about behaviour. We then describe field experiments on clutch size, which show that individuals tend to maximise their lifetime reproductive success. Because an individual’s survival and reproductive success depends critically on its behaviour, selection is expected to design individuals to be efficient at feeding, avoiding predators, finding mates, and so on.

In Chapter 2 we discuss how to test hypotheses for the adaptive advantage of behaviour. One method is comparison among species, in effect analysing the results of evolutionary “experiments”, correlating differences between species in behaviour with differences in ecological and social selection pressures. There have been recent improvements in methodology, using phylogenies to identify independent evolutionary transitions and the order in which traits change. The second method, pioneered by Niko Tinbergen, is to perform experiments, for example to change behaviour and measure the consequences for an individual’s survival and reproductive success.

In Chapter 3, we focus on individual “decision making” between alternative courses of action. We show how optimality models can be used to predict decision rules, and how the same basic models can often be applied to what at first sight seem very different problems, such as feeding and searching for mates. We discuss the roles of social learning and teaching in the development of individual decision making, and use examples from food storing to explore the links between behavioural ecology, cognition and neuroscience. Chapter 4 considers decisions over evolutionary time, and how these change during arms races between predators and prey, and brood parasites and hosts.

The next two chapters consider how individuals should behave when they have to compete with others for scarce resources. Chapter 5 introduces a game theoretic approach to contest behaviour and shows that the outcome is often variability in the population, as individuals distribute across different habitats in space and time, or choose alternative strategies or tactics as they compete for food and mates. We also discuss the concept of animal personalities, a current growing field of research. In Chapter 6, we review the costs and benefits of group living, particularly in relation to foraging and avoiding predation. Recent studies have shown how local decision rules made by individuals can have remarkable consequences for group dynamics, leading to spectacular coordinated movements in bird flocks, fish shoals and ant trails.

The next four chapters are concerned with sexual reproduction. Chapter 7 shows that fundamental differences between the sexes in gamete size and parental investment often leads to males competing for females, either by force or by charm (Darwin’s theory of sexual selection). Females may chose males based on the resources they provide or the genetic benefits for their offspring. There is often sexual conflict and this continues after mating (sperm competition and female choice of sperm). Chapter 8 reviews parental care across the animal kingdom, and three inter-related conflicts: between male and female parent over who should care, and how much care to provide; between siblings; and between parents and their offspring. We consider the theory and evidence for each of these conflicts and distinguish between “battleground” models (which define the conflict) and “resolution” models (which explore the outcomes).

In Chapter 9, we show how different mating systems emerge depending on the economics of parental care and mate defence. The use of DNA profiles to measure parentage has now become routine and has revolutionised our view of family life, revealing (for example) that social monogamy does not necessarily mean genetic monogamy. Chapter 10 examines sex allocation: the problem of how a parent should divide its investment between male and female offspring. Sex allocation in hymenopteran insects provides some of the most convincing quantitative tests of evolutionary theory.

In the final four chapters, we turn our attention to social behaviour. Under what circumstances would we expect the evolution of altruism, namely helping others to reproduce at the expense of one’s own reproductive output? (Chapter 11). When would it pay individuals to cooperate with either related or unrelated individuals? (Chapter 12). We show how social theory can be tested in a variety of animals from microbes to meerkats. Chapter 12 is devoted to the social insects, where altruism reaches its most sophisticated development in the form of sterile worker castes. We discuss new theory for the genetic predispositions and ecological factors promoting this remarkable behaviour, and show that there are often conflicts of interest even within the most cooperative societies. In Chapter 14, we discuss how natural selection shapes signals, focussing on the evolution of honesty and deception.

Finally, in the last chapter (15) we return for a critical re-assessment of some of our main premises: the “gene’s eye” view of behaviour, optimality models and evolutionarily stable strategies. We also point to the flourishing interactions with other fields of research.

The literature has become vast since the earlier editions of this book, so in this edition we have had to be even more selective. We hope that lecturers using this book will add their own favourite examples to those we have mentioned. Throughout, we have tried to point to gaps in current theory and evidence. We hope that readers will be inspired not only to fill these, but also to discover new problems to solve.

Acknowledgements

Photo © Elizabeth Tibbetts.

We thank the editorial staff at Wiley-Blackwell for all their help and encouragement, especially Ward Cooper, Kelvin Matthews and Delia Sandford. Robert Campbell was instrumental in encouraging us in the earlier editions. For help with various chapters, we thank: Joao Alpedrinha, Staffan Andersson, Tim Birkhead, Koos Boomsma, Lucy Browning, Max Burton, Tim Clutton-Brock, Bernie Crespi, Emmett Duffy, Claire El Mouden, Andy Gardner, Ashleigh Griffin, James Higham, Camilla Hinde, Rebecca Kilner, Loeske Kruuk, Carita Lindstedt, Robert Magrath, Allen Moore, Nick Mundy, Hazel Nichols, David Reby, Thom Scott-Phillips, Ben Sheldon, Martin Stevens, Claire Spottiswoode, Mary Caswell (Cassie) Stoddard, Joan Strassmann, Alex Thornton, Rose Thorogood.

And a special thank you to Ann Jeffrey for her truly heroic help with the preparation of the manuscript.

Finally, we thank all our colleagues who so generously provided figures and photographs for the book, and particularly Oliver Krüger for his magnificent cover photograph of a parent Adelie Penguin, introducing its chick to the delights of behavioural ecology.

CHAPTER 1

Natural Selection, Ecology and Behaviour

Photo © Craig Packer

Watching and wondering

Imagine you are watching a bird searching in the grass for food (Fig. 1.1). At first your curiosity may be satisfied simply by knowing what species it is, in this case a starling Sturnus vulgaris. You then watch more closely; the starling walks along and pauses every now and then to probe into the ground. Sometimes it finds a prey item, such as a beetle larva, and eventually, when it has collected several prey items, it flies back to the nest to feed its hungry brood.

Fig. 1.1 A foraging starling. Photo © iStockphoto.com/Dmitry Maslov

For students of behavioural ecology, a whole host of questions comes to mind as this behaviour is observed. The first set of questions concerns how the bird feeds. Why has it chosen this particular place to forage? Why is it alone rather than in a flock? Does it collect every item of food it encounters or is it selective for prey type or size? What influences its decision to stop collecting and fly back to feed its chicks?

Another set of questions emerges when we follow the starling back to its nest. Why has it chosen this site? Why this number of chicks in the nest? How do the two adults decide on how much food each should bring? Are these two adults the mother and father of all the chicks? Why are the chicks begging so noisily and jostling to be fed? Surely this would attract predators to the nest. If we could follow our starlings over a longer period, we may then begin to ask about what determines how much effort the adults put into reproduction versus their own maintenance, about the factors influencing the timing of their seasonal activities, their choice of mate, the dispersal of their offspring and so on.

Asking questions

Behavioural ecology provides a framework for answering these kinds of questions. In this chapter we will show how it combines thinking about behaviour, ecology (the ‘stage’ on which individuals play their behavioural strategies) and evolution (how behaviour evolves by natural selection). But first, we need to be clear about exactly what we mean when we ask the question ‘why?’

Tinbergen’s four ‘why’ questions

Niko Tinbergen (1963), one of the founders of scientific studies of animal behaviour in the wild, emphasized that there are four different ways of answering ‘why’ questions about behaviour. For example, if we asked why male starlings sing in the spring, we could answer as follows:

(1) In terms of causation. Starlings sing because the increasing length of day triggers changes in their hormones, or because of the way air flows through the vocal apparatus and sets up membrane vibrations. These are answers about the mechanisms that cause starlings to sing, including sensory and nervous systems, hormonal mechanisms and skeletal–muscular control.

(2) In terms of development or ontogeny. For example, starlings sing because they have learned the songs from their parents and neighbours, and have a genetic disposition to learn the song of their own species. This answer is concerned with genetic and developmental mechanisms.

(3) In terms of adaptive advantage or function. Starlings sing to attract mates for breeding, and so singing increases the reproductive success of males.

(4) In terms of evolutionary history or phylogeny. This answer would be about how song had evolved in starlings from their avian ancestors. The most primitive living birds make very simple sounds, so it is reasonable to assume that the complex songs of starlings and other song birds have evolved from simpler ancestral calls.

Causal and developmental factors are referred to as proximate because they explain how a given individual comes to behave in a particular way during its lifetime. Factors influencing adaptive advantage and evolution are called ultimate because they explain why and how the individual has evolved the behaviour. To make the distinction clearer, an example is discussed in detail.

Proximate versus ultimate explanations

Reproductive behaviour in lions

In the Serengeti National Park, Tanzania, lions (Panthera leo) live in prides consisting of between three and twelve adult females, from one to six adult males and several cubs (Fig. 1.2a). The group defends a territory in which it hunts for prey, especially gazelle and zebra. Within a pride all the females are related; they are sisters, mothers and daughters, cousins and so on. All were born and reared in the pride and all stay there to breed. Females reproduce from the age of four to eighteen years and so enjoy a long reproductive life.

Fig. 1.2 (a) A lion pride. (i) The females are returning to the middle of their territory after chasing away a neighbouring pride. (ii) Females and cub. (iii) Males patrolling the territory and (iv) relaxing. (v) Male with cub. Photos © Craig Packer (b) Infanticide: a male that has just taken over ownership of a pride, with a cub in his jaws that he has killed. Photo © Tim Caro

For the males, life is very different. When they are three years old, young related males (sometimes brothers) leave their natal pride. After a couple of years as nomads they attempt to take over another pride from old and weak males. After a successful takeover they stay in the pride for two to three years before they, in turn, are driven out by new males. A male’s reproductive life is therefore short.