Cover

Half-title

Title

Copyright

Dedication

Contents

List of Contributors

Preface

1 The Study of Mammalian Cooperative Breeding

1.1 Introduction

1.2 Terminology

1.3 The book

1.4 Conclusion

References

2 The Bioenergetics of Parental Behavior and the Evolution of Alloparental Care in Marmosets and Tamarins

2.1 Alloparental behavior

2.2 Social organization and infant care in callitrichids

2.3 Potential benefits of alloparental behavior

2.3.1 Hypotheses

2.3.1.1 Benefits from experience

2.3.1.2 Benefit through inheritance of a breeding position

2.3.1.3 Benefit of infant care as courtship

2.3.1.4 Benefit through improved indirect fitness

2.4 Costs of alloparental behavior

2.4.1 Levels of cost

2.4.1.1 Immediate costs of alloparental behavior

2.5 Evolution of alloparental behavior in marmosets and tamarins

2.6 Conclusions

Acknowledgments

References

3 Proximate Regulation of Singular Breeding in Callitrichid Primates

3.1 Introduction

3.2 Proximate mechanisms maintaining singular breeding in callitrichid primates

3.2.1 Physiological mechanisms

3.2.1.1 Puberty delay

3.2.1.2 Suppression of ovulation

3.2.1.3 Postconception physiological regulation of fertility in subordinates

3.2.2 Behavioral mechanisms maintaining singular breeding

3.2.2.1 Preconception mechanisms

3.2.2.2 Postconception mechanisms

3.3 Plural breeding in social groups of callitrichid primates

3.3.1 Departures from singular breeding in wild populations

3.3.1.1 Pygmy marmosets (genus Cebuella,)

3.3.1.2 Marmosets (genus Callithrix,)

3.3.1.3 Tamarins (genus Saguinusj

3.3.1.4 Lion tamarins (genus Leontopithecusj

3.3.1.5 Goeldi's monkey (genus callimico,)

3.3.2 Departures from singular breeding in captive populations

3.4 Species differences in mechanisms of reproductive suppression: the cost of reproduction

3.5 Conclusions

Acknowledgments

References

4 Cooperative Breeding, Reproductive Suppression, and Body Mass in Canids

4.1 Introduction

4.2 Methods

4.2.1 Data selection

4.2.2 Statistical analysis

4.3 Results

4.3.1 Allometry of life-history traits

4.3.1.1 Group size and feeding ecology

4.3.1.2 Cooperative hunting

4.3.1.3 Parental and alloparental input

4.3.1.4 Pup survival

4.3.2 Modeling parental and alloparental input

4.3.2.1 The allometry of parental input

4.3.2.2 The consequences of alloparental input

4.3.2.3 Reproductive suppression

4.3.2.4 Resources, philopatry, and dispersal

4.4 Discussion

4.5 Conclusions

Appendix

Acknowledgments

References

5 Hormonal and Experiential Factors in the Expression of Social and Parental Behavior in Canids

5.1 Introduction

5.2 The basic canid social system

5.3 Social aspects of reproduction

5.3.1 Paternal care

5.3.2 Incorporation of postpubertal young

5.3.3 Reproductive suppression

5.4 Reproductive physiology

5.4.1 Seasonal reproduction

5.4.1.1 Monestrum and the female cycle

5.4.1.2 Male seasonality

5.4.1.3 Prolactin

5.5 Phases of the ovulatory cycle

5.5.1 The anestrous phase

5.5.2 Proestrous phase

5.5.3 Estrous phase

5.5.4 Pregnancy

5.5.5 Luteal phase or pseudopregnancy

5.6 The endocrine basis of parental behavior

5.7 The interplay of physiology and social organization

5.8 Conclusions

References

6 Variation in Reproductive Suppression among Dwarf Mongooses: Interplay between Mechanisms and Evolution

6.1 Introduction

6.1.1 Why suppress subordinates?

6.1.2 Why accept suppression?

6.1.3 Variation in the degree of suppression

6.2 Dwarf mongoose social organization

6.2.1 The study population

6.2.2 Social organization

6.3 Evolutionary modeling of subordinate reproduction

6.4 Evolutionary predictions for subordinate reproduction

6.4.1 Estimating parameter values

6.4.1.1 Relatedness

6.4.1.2 Effects of subordinates on breeders9 reproduction

6.4.1.3 Dispersal risk

6.4.1.4 Probabilities of obtaining an alpha position

6.4.1.5 Postdispersal reproductive success

6.4.2 Predicted patterns of subordinate reproduction

6.4.3 Observed patterns of subordinate reproduction

6.4.3.1 Mean degree of suppression

6.4.4 Mechanistic influences on patterns of subordinate reproduction

6.5 Summary

Acknowledgments

References

7 Dynamic Optimization and Cooperative Breeding: An Evaluation of Future Fitness Effects

7.1 Introduction

7.2 The animals

7.3 The model

7.3.1 The dynamic program

7.4 Forward iteration

7.5 Decisions based on immediate payoffs

7.6 Lifetime inclusive fitness, fitness gradients, and parameter estimates

7.7 Results

7.8 Future effects

7.9 Indirect effects

7.10 Discussion

7.11 Conclusions

Acknowledgments

References

8 Examination of Alternative Hypotheses for Cooperative Breeding in Rodents

8.1 Introduction

8.2 Identification of cooperative breeding in rodents

8.3 Evolution and maintenance of philopatry

8.4 Evolution and maintenance of reproductive suppression

8.5 Evolution and maintenance of alloparental behavior

8.6 Conclusions

Acknowledgments

References

9 The Psychobiological Basis of Cooperative Breeding in Rodents

9.1 Introduction

9.2 Monogamy and cooperative breeding

9.3 Social stimuli and female reproduction

9.3.1 Behavioral requirements for reproductive activation in female prairie voles

9.3.2 Neuroendocrine correlates of reproductive activation in female prairie voles

9.3.3 Possible functions of extended copulatory interactions

9.3.4 Postpartum estrus

9.3.5 Reproductive inhibition by social factors

9.3.5.1 Inhibition of estrus induction

9.3.5.2 Incest avoidance

9.4 Social stimuli and male reproductive behavior

9.4.1 Male reproductive activation

9.4.2 Social suppression of male reproduction

9.4.3 Factors regulating patterns of male sexual behavior

9.5 Pair bonding

9.5.1 Physiology of partner preference formation in female prairie voles

9.5.2 Physiology of partner preferences in male prairie voles

9.5.3 Physiology of selective aggression in prairie voles

9.5.4 Summary of factors regulating pair bonding

9.6 Proximate factors associated with alloparenting and philopatry

9.6.1 Proximate factors associated with parental behavior

9.6.2 Prior experience and alloparental behavior

9.6.3 Pheromonal factors and parental behavior

9.6.4 Organizational factors and parental behavior

9.6.5 Organizational factors and philopatry

9.6.6 Population differences in the expression of alloparenting

9.6.7 Summary of proximate mechanisms responsible for alloparenting

9.7 Summary

Acknowledgments

References

10 Cooperative Breeding in Naked Mole-Rats: Implications for Vertebrate and Invertebrate Sociality

10.1 Introduction

10.2 Natural history of naked mole-rats

10.3 Social organization of naked mole-rats

10.3.1 Coloniality

10.3.1.1 Naked mole-rats live in groups

10.3.1.2 Naked mole-rat colonies are extended family groups

10.3.1.3 New colony formation is poorly understood

10.3.2 Reproductive division of labor

10.4 Cooperative care of young

10.5 Colony regulation

10.6 Evolution of cooperative breeding in naked mole-rats

10.7 Implications for vertebrate and invertebrate sociality

10.8 Sociality in other subterranean rodents

10.9 Summary

Acknowledgments

References

11 The Physiology of a Reproductive Dictatorship: Regulation of Male and Female Reproduction by a Single Breeding Female in Colonies of Naked Mole-Rats

11.1 Introduction

11.2 Environmental regulation of mammalian reproduction

11.2.1 Social status and reproductive success

11.3 Cooperatively breeding species

11.4 African mole-rats and the occurrence of cooperative breeding and reproductive suppression

11.5 Naked mole-rats

11.5.1 Sexual and agonistic interactions in captive colonies

11.5.2 Physiology of reproductive suppression in nonbreeding males and females

11.5.3 Regulation of reproductive physiology in breeding males

11.5.4 Cues regulating male and female reproduction

11.5.5 Relevance of reproductive adaptations to cooperative breeding

11.6 Conclusions

Acknowledgments

References

12 Factors Influencing the Occurrence of Communal Care in Plural Breeding Mammals

12.1 Introduction

12.2 Definitions and ultimate explanations for communal care

12.3 Why live and breed in groups?

12.4 Types of alloparental care

12.4.1 Formation of nursery groups for the benefit of dilution

12.4.2 Formation of nursery groups for the benefit of thermoregulation

12.4.3 Communal defense of young

12.4.4 "Babysitting"

12.4.5 Adoption

12.4.6 Group provisioning

12.4.7 Helper-assisted birth

12.4.8 Nonoffspring nursing

12.5 Comparative study of nonoffspring nursing

12.6 Comparison of nonoffspring nursing in lions and hyenas

12.7 Factors influencing variation in nonoffspring nursing in lions

12.8 Conclusions

12.9 Directions for future research

Acknowledgments

References

13 A Bird's-Eye View of Mammalian Cooperative Breeding

13.1 Introduction

13.2 Fundamental biological differences between birds and mammals

13.3 Why do groups form?

13.3.1 Within-population variation in delayed dispersal

13.3.2 Among-population variation in delayed dispersal

13.3.3 Interspecific variation in delayed dispersal

13.4 What determines who breeds?

13.4.1 Variation in breeding suppression: proximate causes

13.4.2 Variation in breeding suppression: ultimate causes

13.5 Why do some individuals provide alloparental care?

13.5.1 Alloparental care: proximate causes

13.5.2 Alloparental care: ultimate causes

13.5.2.1 Adaptive hypotheses based on future direct fitness

13.5.2.2 Adaptive hypotheses based on present indirect fitness

13.5.2.3 Adaptive hypotheses based on future indirect fitness

13.5.2.4 Nonadaptive hypotheses for alloparental behavior

13.5.2.5 Costs of alloparental care

13.6 Cooperative breeding in mammals: future directions

13.7 Conclusion: toward "the new synthesis"

Acknowledgments

References

Index