Living in Groups

Group Living is a widespread phenomenon within the animal kingdom and has attracted considerable attention in a number of different contexts. This book is focused on the unifying concepts regarding group behaviour that have been developed over the last two decades.The authors set out to discuss the mechanisms that govern the evolution and maintenance of grouping behaviour throughout the animal kingdom, and the ecological factors that control observed group size and group composition in particular situations. Although the book's emphasis is on the elaborationof a conceptual framework, extensive examples and case studies illustrate the diversity of grouping phemonema across taxonomic boundaries, and demonstrate the general applicability of the concepts involved.This book will familiarise the reader with the latest ideas on the ecology and evolution of group-living animals, providing a summary and critical synthesis of the extensive and diverse literature on the subject.

Introduction

1

(5)

Overview

1

(1)

A definition of groups

2

(2)

Book structure

4

(2)

The benefits of group formation

6

(35)

Introduction

6

(2)

Anti-predator vigilance

8

(5)

The classical many-eyes theory

8

(1)

How individual vigilance works

9

(1)

Information transfer between individuals

10

(2)

Some unanswered questions on group vigilance

12

(1)

Related issues

12

(1)

Dilution of risk

13

(6)

Avoidance, dilution, and abatement

13

(4)

Predator swamping

17

(1)

The Selfish herd

17

(1)

Defence against parasites

17

(2)

Preadator confusion

19

(4)

Theory

19

(1)

Empirical support for theoretical predictions

20

(1)

Cognitive limitations

21

(1)

Communal defence against predators

22

(1)

Predator learning

23

(1)

Foraging benefits to grouping

23

(9)

Benefits for predators

23

(2)

Finding food

25

(7)

Finding a mate

32

(3)

Conserving heat and water

35

(2)

Reducing the energetic costs of movement

37

(3)

Introduction

37

(1)

Movement in water

37

(2)

Movement in air

39

(1)

Summary and conclusions

40

(1)

Some costs to grouping

41

(14)

Introduction

41

(1)

Increased attack rate on larger groups

42

(4)

Foraging in a group

46

(5)

Kleptoparasitism

47

(1)

Aggression more generally

47

(1)

Pseudo-interference

48

(1)

Shadow interference of sit and wait predators

48

(1)

Just getting in each other's way

49

(1)

Prey response to detected predators

50

(1)

A note on generality

51

(1)

Increased parasite burdens

51

(1)

Misdirected parental care

52

(2)

Cuckoldry

52

(1)

Brood parasitism and adoption

53

(1)

Summary and conclusions

54

(1)

The size of a group

55

(18)

Introduction

55

(3)

Combining costs and benefits of grouping

55

(1)

An illustrative example

56

(1)

The shape of the fitness function

56

(2)

Are optimal group sizes likely to be seen in nature?

58

(10)

An argument why groups should be larger than optimal

58

(2)

Refinements of the argument

60

(1)

The role of relatedness

61

(1)

The influence of competition

62

(1)

The effect of dominance hierarchies

63

(1)

Empirical evidence for active recruitment to foraging groups

63

(5)

Observed group sizes in nature

68

(3)

Social carnivores

68

(1)

Data from other taxa

69

(1)

Distribution of group sizes

69

(2)

Summary and conclusions

71

(2)

Spatial heterogeneity of costs and benefits within groups

73

(14)

Introduction

73

(1)

Group structure and spatial positions: definitions

74

(1)

Cause and effect relationships

75

(2)

Energy gains and losses

77

(3)

Energy intake

77

(1)

Energy expenditure

78

(1)

Net-energy payoff

79

(1)

Predation risk

80

(3)

Stationary groups

80

(1)

Mobile groups

80

(3)

Parasites

83

(1)

Reproductive success

83

(1)

Dominance status

84

(1)

Trade-offs between different fitness currencies

85

(1)

Summary and conclusions

86

(1)

Heterogeneity and homogeneity of group membership

87

(17)

Introduction

87

(1)

Theory of assortativeness

88

(1)

The influence of predation on assortment

89

(8)

Predator preference for odd prey (the oddity effect)

90

(3)

Evidence for the oddity effect from prey behaviour

93

(1)

Predator preference for common prey

93

(1)

Frequency-independent preferences

94

(2)

Reduced vigilance in mixed species groups

96

(1)

Differential anti-predatory abilities within a group

96

(1)

Evidence for the evolution of group mate preferences in prey

97

(4)

The role of species

97

(1)

The role of body length and colour

98

(1)

The role of parasitism

99

(1)

The role of familiarity

100

(1)

The role of kinship

100

(1)

Multi-species foraging groups

101

(1)

Consequences of inter-individual differences for optimal group size

102

(1)

Summary and conclusions

102

(2)

Evolutionary considerations

104

(19)

Introduction

104

(1)

Individual differences: artificial selection

104

(2)

Population differences

106

(4)

Testing for population differences

106

(4)

Species differences

110

(9)

Pathways towards the evolution of groups

113

(6)

Groups as units of selection

119

(2)

Summary and conclusions

121

(2)

Environmental effects on grouping behaviour

123

(14)

Introduction

123

(1)

Ontogenetic constraints on grouping: fish shoaling behaviour

123

(2)

Ontogenetic shift in sociality: the spiny lobster

125

(1)

The role of rearing conditions: caste determination and division of labour in the honey bee

126

(2)

Behavioural changes induced by crowding: the desert locust

128

(2)

The role of learning

130

(2)

Parasite-mediated changes in behaviour

132

(3)

Summary and conclusions

135

(2)

Mechanisms

137

(14)

Introduction

137

(1)

Recognition of suitable group mates

137

(1)

Inter-individual distance regulation: attraction and repulsion

138

(2)

Group formation

140

(1)

Collective behaviour

140

(6)

Locomotion

140

(2)

Positioning behaviour and leadership

142

(1)

Group structure

143

(2)

Collective foraging

145

(1)

Teams

146

(1)

Assessment of group size and group composition

146

(1)

Group size distributions

147

(3)

Summary and conclusions

150

(1)

Conclusions

151

(8)

Introduction

151

(1)

Group size

151

(4)

Individual-based models of group size

152

(1)

Population density and group size

153

(2)

Comparative studies

155

(1)

Evolution of grouping

156

(1)

Group composition

156

(1)

Signalling

157

(1)

Short-term behavioural change

157

(1)

Concluding remarks

158

(1)

References

159

(30)

Author Index

189

(10)

General Index

199

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Summary

Table of Contents

Supplemental Materials