Essays in Animal Behaviour: Celebrating 50 Years of Animal Behaviour

575 Pages·2005·23.98 MB·English

by  Jeffrey R Lucas

Marine Mammals:
Evolutionary Biology
2nd Edition
This Page Intentionally Left Blank
Marine Mammals:
Evolutionary Biology
2nd Edition
Annalisa Berta
Department ofBiology
San Diego State University
San Diego,California
James L.Sumich
Biology Department
Grossmont College
El Cajon,California
Kit M.Kovacs
Biodiversity Programme
Norwegian Polar Institute
Tromsø,Norway
With illustrations by
Pieter Arend Folkens
A Higher Porpoise Design Group
Benecia,California
Peter J.Adam
Department ofBiology
San Diego State University
San Diego,California
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ISBN 13:978-0-12-369499-7
ISBN 10:0-12-088552-2
Printed in the United States ofAmerica
05 06 07 08 09 10 9 8 7 6 5 4 3 2 1
CONTENTS
Preface ix
Acknowledgments x
1. Introduction 1
1.1. Marine Mammals—“What Are They?” 1
1.2. Adaptations for Aquatic Life 1
1.3. Scope and Use ofThis Book 2
1.4. Time Scale 2
1.5. Early Observations ofMarine Mammals 3
1.6. Emergence ofMarine Mammal Science 7
1.7. Further Reading and Resources 9
References 9
PARTI: EVOLUTIONARY HISTORY
2. Systematics and Classification 12
2.1. Introduction:Systematics—What Is It and Why Do It? 12
2.2. Some Basic Terminology and Concepts 13
2.3. How Do You Do Cladistics? 17
2.4. Testing Phylogenetic Hypotheses 19
2.5. Going Beyond the Phylogenetic Framework:Elucidating
Evolutionary and Ecological Patterns 21
2.6. Taxonomy and Classification 22
2.7. Summary and Conclusions 24
2.8. Further Reading 24
References 24
v
vi Contents
3. Pinniped Evolution and Systematics 27
3.1. Introduction 27
3.2. Origin and Evolution 27
3.3. Summary and Conclusions 46
3.4. Further Reading 47
References 47
4. Cetacean Evolution and Systematics 51
4.1. Introduction 51
4.2. Origin and Evolution 51
4.3. Summary and Conclusions 81
4.4. Further Reading 81
References 82
5. Sirenians and Other Marine Mammals:Evolution and Systematics 89
5.1. Introduction 89
5.2. Origin and Evolution ofSirenians 89
5.3. The Extinct Sirenian Relatives—Desmostylia 98
5.4. The Extinct Marine Bear-Like Carnivoran,Kolponomos 100
5.5. The Extinct Aquatic Sloth,Thalassocnus natans 102
5.6. The Sea Otter,Enhydra lutris 102
5.7. The Polar Bear,Ursus maritimus 105
5.8. Summary and Conclusions 105
5.9. Further Reading 107
References 107
6. Evolutionary Biogeography 111
6.1. Introduction—What Is Biogeography and Why Is It Important? 111
6.2. Ecological Factors Affecting Distributions ofMarine Mammals 111
6.3. Present Patterns ofDistribution 117
6.4. Reconstructing Biogeographic Patterns 119
6.5. Past Patterns ofDistribution 121
6.6. Summary and Conclusions 127
6.7. Further Reading and Resources 128
References 129
PARTII:EVOLUTIONARYBIOLOGY,ECOLOGY,ANDBEHAVIOR
7. Integumentary and Sensory Systems 132
7.1. Introduction 132
7.2. Integumentary System 132
7.3. Nerves and Sense Organs 148
7.4. Summary and Conclusions 156
Contents vii
7.5. Further Reading 157
References 157
8. Musculoskeletal System and Locomotion 165
8.1. Introduction 165
8.2. Pinnipeds 165
8.3. Cetaceans 178
8.4. Sirenians 195
8.5. Sea Otter 200
8.6. Polar Bear 203
8.7. Summary and Conclusions 203
8.8. Further Reading 205
References 206
9. Energetics 213
9.1. Introduction 213
9.2. Metabolic Rates 213
9.3. Thermoregulation 217
9.4. Energetics ofLocomotion 223
9.5. Osmoregulation 229
9.6. Summary and Conclusions 231
9.7. Further Reading 232
References 232
10. Respiration and Diving Physiology 237
10.1. Introduction 237
10.2. Problems ofDeep and Prolonged Dives for Breath-Holders 237
10.3. Pulmonary and Circulatory Adaptations to Diving 239
10.4. Diving Response 252
10.5. Diving Behavior and Phylogenetic Patterns 254
10.6. Summary and Conclusions 262
10.7. Further Reading 262
References 262
11. Sound Production for Communication,Echolocation,and Prey Capture 270
11.1. Introduction 270
11.2. Sound Propagation in Air and Water 270
11.3. Anatomy and Physiology ofSound Production and Reception 272
11.4. Functions ofIntentionally Produced Sounds 284
11.5. Acoustic Thermometry ofOcean Climate and Low-Frequency
Military Sonars 302
11.6. Summary and Conclusions 303
11.7. Further Reading 304
References 305
viii Contents
12. Diet,Foraging Structures,and Strategies 312
12.1. Introduction 312
12.2. Seasonal and Geographical Patterns ofPrey Abundance 313
12.3. Adaptations for Foraging in Pinnipeds 314
12.4. Feeding Specializations ofCetaceans 324
12.5. Feeding Specializations ofSirenians 344
12.6. Feeding Specializations ofOther Marine Mammals 350
12.7. Summary and Conclusions 354
12.8. Further Reading 355
References 355
13. Reproductive Structures,Strategies,and Patterns 363
13.1. Introduction 363
13.2. Anatomy and Physiology ofthe Reproductive System 365
13.3. Mating Systems 377
13.4. Lactation Strategies 395
13.5. Reproductive Patterns 402
13.6. Summary and Conclusions 405
13.7. Further Reading 406
References 407
14. Population Structure and Dynamics 416
14.1. Introduction 416
14.2. Abundance and Its Determination in Marine Mammals 417
14.3. Techniques for Monitoring Populations 419
14.4. Population Structure and Dynamics 432
14.5. Summary and Conclusions 444
14.6. Further Reading 445
References 445
15. Exploitation and Conservation 456
15.1. Introduction 456
15.2. Commercial Exploitation ofMarine Mammals 456
15.3. Legal Framework for Marine Mammal Conservation and Protection 460
15.4. Incidental Taking ofMarine Mammals 466
15.5. Environmental Contaminants 472
15.6. Single Beachings vs Mass Strandings 475
15.7. Ecotourism 478
15.8. Progress and the Future 480
15.9. Summary and Conclusions 483
15.10. Further Reading 483
References 484
Appendix:Classification ofMarine Mammals 491
Glossary 513
Index 521
PREFACE
The second edition,like the previous one,Marine Mammals:Evolutionary Biology,is
written for two audiences:as a text for an upper-level undergraduate or graduate-level
course on marine mammal biology and as a source book for marine mammal scientists
in research, education,management,and legal/policy development positions.One of
our major goals is to introduce the reader to the tremendous breadth oftopics that com-
prise the rapidly expanding interdisciplinary field ofmarine mammal science today.Our
motivation for writing this book was the lack ofa comprehensive text on marine mam-
mal biology,particularly one that employs a comparative,phylogenetic approach.We
have attempted,where possible,to demonstrate that hypotheses ofthe evolutionary rela-
tionships ofmarine mammals provide a powerful framework for tracing the evolution of
their morphology,behavior,and ecology.This approach has much to offer but is limited,
in many cases,by available comparative data.We hope that this book stimulates others
to pursue marine mammal research in this exciting new direction.
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