Ethology and Behavioral Ecology of Otariids and the Odobenid

Description

This book is focused on the marine mammalian groups the Otariidae and the Odobenidae, otherwise known as fur seals, sea lions and the walrus. In 30 chapters, more than 60 authors from 30 institutions and 13 nationalities, discuss a broad suite of topics from maternal care and mating behavior, through play, cognition and personality, to adaptation to life in the Anthropocene. The authors explore the behaviors that have allowed these semi-aquatic mammals to thrive in the marine realm. Many populations have recovered following historical decimation, with interesting evolutionary consequences which are explored. Detailed, selected, individual species descriptions are also provided, showcasing the behavioral diversity of this engaging, adaptive and highly successful group of marine mammals. Introduction to the Series Preface Contents Part I: Behavior on Land and at Sea: Introduction Chapter 1: Otariid Ethology: One Researcher´s Historical Perspective 1.1 Introduction 1.2 Otariid Study Methods Over Time 1.3 Otariid Life Cycles 1.4 Otariid Annual Cycles 1.5 Social Behavior and Organization 1.6 Otariids and the Society for Marine Mammalogy 1.7 Manipulative Behavioral Experiments 1.8 Research on Diving Behavior 1.9 Research Using Isotopes 1.10 Concluding Remarks References Chapter 2: When Physiology and Ecology Meet: The Interdependency Between Foraging Ecology and Reproduction in Otariids 2.1 Introduction 2.2 Energetics of Growth, Reproduction and an Amphibious Lifestyle 2.2.1 Tradeoffs in Time on Land and at Sea 2.2.2 Diving Physiology and Capacity 2.2.3 Field Metabolic Rates 2.3 Habitat Diversity, Foraging Patterns and Reproductive Strategies 2.3.1 Foraging Behavior Relative to Habitats and Energetics 2.3.2 Consequences for Population Dynamics 2.3.3 Role of Food Availability 2.4 Evolutionary Implications of Otariid Foraging Patterns 2.5 Future Directions References Chapter 3: Maternal Behavior in Otariids and the Walrus 3.1 Introduction 3.2 Gestation Length and Maternal Attendance Patterns 3.3 Parental Care 3.4 Mother-Offspring Conflicts and Weaning 3.5 Conclusions References Part II: Foraging Behavior Chapter 4: Foraging Capacities, Behaviors and Strategies of Otariids and Odobenids 4.1 Introduction 4.2 Morphological and Physiological Diving Capacities of Fur Seals and Sea Lions 4.3 Diving Strategies of Fur Seals and Sea Lions 4.4 Dive Metrics to Infer Behaviors and Energetics 4.5 Conclusion: Impacts of Diving Strategies on Fitness and Population Trends References Chapter 5: Prey Capture and Processing in Fur Seals, Sea Lions and the Walrus 5.1 Introduction 5.1.1 Challenges of Feeding in Water 5.1.2 Origin and Structure of Otarioid Feeding Anatomy 5.2 The Feeding Cycle 5.3 Feeding in Otariids 5.3.1 Diet 5.3.2 Semi-Aquatic and Raptorial Feeding 5.3.3 Suction Feeding 5.4 Feeding in Walruses 5.4.1 Diet 5.4.2 Suction Feeding 5.4.3 Feeding on Large Prey 5.5 Conclusions References Part III: Reproductive Behavior Chapter 6: Polygyny in the Era of Molecular Biology: Revisiting Bartholomew´s Model 6.1 Introduction 6.2 Methods 6.3 Summary of Results from Paternity Studies 6.4 Does Theory Meet Facts? 6.5 A Mixed Model of the Evolution of Sexual Size Dimorphism and Mating Systems in Mammals 6.6 Final Remarks References Chapter 7: The Role of Females in the Evolution of Sexual Dimorphism and Mating Systems 7.1 Introduction 7.2 Evolution of Body Size in Pinnipeds 7.3 Role of Females in the Evolution of Sexual Size Dimorphism: Fecundity Selection 7.4 Role of Females in the Evolution of Body Size in Pinnipeds: Lactation Strategies 7.5 Role of Females in the Evolution of Mating Systems: Avoidance of Male Harassment 7.6 Role of Females in Sexual Selection: Extra-Harem Paternity, Lekking and Mate Choice 7.7 Conclusions References Chapter 8: Site Fidelity, Male Harassment and Female Gregariousness: Factors Shaping the Highly Polygynous Mating System of th... 8.1 Introduction 8.2 Longitudinal Behavioral Study 8.3 Phenology of Reproductive Behavior 8.3.1 Natal Philopatry of Males 8.4 Territory Acquisition 8.5 How Males Find Mating Partners 8.6 Male-Female Association and Paternity 8.7 Kleptogyny and Other Alternative Strategies of Less Successful Males 8.8 Harassment of Females and Pups by Non-territorial Males 8.9 Female Choice of Breeding Sites 8.10 Extreme Polygyny References Chapter 9: Mating Games and Raiding Parties in Southern Sea Lions 9.1 Genesis 9.2 Objectives 9.3 The Mating System of the 1980s 9.4 Group Raids and Its Consequences 9.5 Group Raids from the Male and Female Perspective 9.6 Mating Systems in Other Places 9.7 Mating System at Punta Norte in 2020 9.8 Female Safety in 2020 9.9 Other Species 9.9.1 Human Raids 9.10 General Conclusions References Chapter 10: Infanticide in Eared Seals 10.1 Intentional Infanticide 10.1.1 Intra-Specific 10.1.2 Inter-Specific 10.2 Unintentional Infanticide 10.2.1 By-Product of Sexual Behavior and Intra-Sexual Competition 10.2.2 By-Product of Misdirected Aggression 10.2.3 By-Product of Adoption Avoidance Behavior 10.2.4 By-Product of Competition for Mis-recognised Pups 10.2.5 By-Product of Rough Handling by Mothers 10.2.6 Incidental Trampling and Crushing 10.3 Conclusions References Chapter 11: Genetic Consequences of Dispersal, Philopatry and Reproductive Behaviors 11.1 Introduction 11.2 Male Harassment as a Central Driver of Otariid Gregariousness 11.3 Gregariousness and the Metapopulation Model 11.4 Natal Philopatry and Breeding Site Fidelity 11.4.1 The Consequences of Dispersal on Genetic Variation 11.4.2 Fixing Genetic Differences: Evolution at Work 11.4.3 Overcoming Natal Philopatry: Otariid Recolonization and Its Nuances 11.4.3.1 Implications for Management 11.5 Polygyny 11.5.1 The Many Shades of Polygyny 11.6 Otariid Behavior and Sealing 11.6.1 The Long-Term Consequences of Sealing: What Have We Learned? 11.7 Living in a Dynamic Environment: The Walrus References Chapter 12: Intergeneric Fertile Hybridization in Otariids References Chapter 13: Visual Species Recognition and Mate Choice in Fur Seals: An Experimental Study 13.1 Introduction 13.2 Materials and Methods 13.3 Analysis 13.4 Results 13.5 Discussion References Part IV: Cognitive Behavior, Communication, Personality and Behavioral Development Chapter 14: Vocal Communication in Otariids and Odobenids 14.1 Introduction 14.1.1 Vocal Communication 14.1.2 Vocal Repertoire and Vocalizations 14.1.3 Information and Functions of Vocalizations 14.2 Vocal Communication in Otariids and Odobenids 14.3 Mother-Pup Vocal Communication 14.3.1 Vocal Repertoire 14.3.1.1 In Otariids 14.3.1.2 In Odobenid 14.3.2 Vocal Stereotypy and Evidence for Individual Recognition 14.3.3 Ontogeny of Vocal Recognition 14.3.4 Vocal Development and Long-Term Recognition 14.3.5 Individual Vocal Signature 14.4 Vocal Communication in Males 14.4.1 The Vocal Repertoire 14.4.1.1 Otariids 14.4.1.2 In Odobenids 14.4.2 Vocal Recognition and/or Rival Assessment 14.4.2.1 In Otariids 14.4.2.2 In Odobenids 14.5 Conclusions References Chapter 15: Non-vocal Communication in Otariids and Odobenids: The Involvement of Visual and Olfactory Cues in Their Social Li... 15.1 Introduction 15.1.1 Visual Cues 15.1.2 Olfactory Cues 15.2 Visual Cues Involved in Social Recognition in Otariids and Odobenids 15.2.1 Visual Abilities 15.2.2 Visual Cues in Mother-Offspring Relationship 15.2.3 Visual Cues Used in Male-Male Interactions 15.3 Olfactory Cues Involved in Social Recognition in Otariids and Odobenids 15.3.1 Olfactory Abilities 15.3.2 Olfactory Cues in Mother-Pup Interactions 15.3.3 Olfactory Cues Used in Male-Male Interactions 15.4 Conclusions References Chapter 16: The Sensory World of Otariids 16.1 Introduction 16.2 The Visual System 16.3 Mechanoreception 16.3.1 Haptics and Hydrodynamics 16.3.2 Audition 16.4 Chemoreception 16.5 Other Senses 16.6 Conclusions References Chapter 17: The Mind of a Sea Lion 17.1 Introduction 17.2 Foundational Research 17.3 Concept Formation 17.4 Learning in the Vocal Domain 17.5 Rhythmic Behavior 17.6 Temperament and Self-Regulation 17.7 Sea Lions as Natural Models for Studying Brain and Behavior 17.8 Conclusions References Chapter 18: The California Sea Lion: Thriving in a Human-Dominated World 18.1 Introduction 18.2 Learning in a Changing World 18.3 Single Stimulus Learning: Habituation and Sensitization in a Changing World 18.4 Management Implications of Habituation/Sensitization 18.5 Associative Learning via Pavlovian and Instrumental Conditioning 18.6 Management Implications of Associative Learning 18.7 A Learning-Based Approach to Deterrents 18.8 Social Learning Mechanisms 18.9 Management Implications of Social Learning 18.10 Conclusions References Chapter 19: Pinniped `Personality´, or Consistent Individual Differences 19.1 Introduction 19.2 Recognizing Animal Personalities 19.2.1 Historic Perspectives 19.2.2 Ecological Applications 19.2.3 Modern Methodologies 19.3 Capturing Individual Differences in Pinnipeds 19.4 Ontogeny of Pup Personality 19.4.1 Internal Mechanisms and External Stimuli 19.4.2 Early Life Challenges 19.4.3 Social Interactions and Play 19.4.4 Maternal Effects 19.5 Reproductive Consequences 19.5.1 Life History Theory 19.5.2 Fortune Favors the Bold? 19.6 Individual Foraging Strategies 19.6.1 Drivers of Intraspecific Variation 19.6.2 Consistency: The Key to Success? 19.7 Implications for Conservation 19.8 Final Questions References Chapter 20: Playing at the Edge of the Sea: A Comparative Analysis in Otariids and Odobenids 20.1 Introduction 20.2 One Step Before We `Playfully´ Jump into Our Target Group 20.3 Different Maternal Styles Matter: The Ontogenetic Pathways of Social Play 20.3.1 Looking at the Mother: Mother-Pup Play 20.3.2 Looking at Peers: Pup-Pup Play 20.4 Towards Maternal Independence: Play Among Juveniles 20.5 To Be Predator or Prey: The Role of Solitary Play in Developing Future Successful Strategies 20.6 Conclusion References Part V: Diving Deeper into Species Chapter 21: Responding to Human Influence: Southern Sea Lion Males Adapt to Harbor Habitats 21.1 Introduction 21.2 Origin of the Harbor Colonies 21.3 Age Structure 21.4 Long Term Trends in Abundance 21.5 Habitat Use and Internal Distribution 21.6 Food Provisioning and Foraging 21.7 Functional Connections with Breeding Colonies 21.8 Social Structure and Individual Association 21.9 Activity Budgets and Agonistic Interactions 21.10 Management Challenges in Harbor Habitats 21.11 Conclusions References Chapter 22: Ethology and Behavioral Ecology of the Walrus (Odobenus rosmarus), with Emphasis on Communication and Social Behav... 22.1 Introduction 22.2 Walrus Tusks as Social Organs 22.3 Notes on Behavioral Description 22.4 Walrus Breeding Behavior and Male Displays 22.5 Optical and Tactile Communication 22.6 Gregariousness and Thigmotaxis 22.7 Herds and Terrestrial Haulouts 22.8 Social System 22.9 Movements and Diving: Descriptions Across Scales 22.10 Feeding 22.11 Concluding Comments References Chapter 23: Behavioral Insights into the Decline and Natural History of Steller Sea Lions 23.1 Introduction 23.2 Natural History 23.3 Breeding Behavior 23.4 Behavioral Proxies of Environmental Conditions 23.5 Weaning Behavior 23.6 Abortions and Stillbirths 23.7 Conclusions References Chapter 24: Exploitation and Recovery of the South American Sea Lion in the Southwestern Atlantic 24.1 Introduction 24.2 History of Exploitation 24.2.1 Pre-commercial Period: 7000 BP Until the Colonization of America 24.2.2 Colonization by Europeans 24.3 History of Recovery 24.3.1 Demographics 24.3.2 New Colonies 24.3.3 Female Behavior 24.4 Conclusions: Old Problems-New Problems 24.5 Future Scenarios References Chapter 25: Living on the Edge, the New Zealand Sea Lion 25.1 Introduction 25.2 Background 25.2.1 Distribution 25.2.2 Population Trends and Conservation Status 25.3 Site Fidelity, Philopatry and Recolonization Behavior 25.4 Restless Maternal Behavior 25.4.1 Female Movement and Male Harassment: Dimorphism and Polygamous Breeding 25.4.2 Female Movement, Parasites and Disease 25.4.3 Female Movement and Foraging Behavior 25.5 Conclusions References Chapter 26: The Enigmatic Life History of the Australian Sea Lion 26.1 Genetics and Biogeography 26.2 Conservation Status 26.3 Life History 26.3.1 Breeding Cycle and Demography 26.3.2 Diet and Foraging Behavior: Near Apex Predator 26.4 Parent-Offspring Behavior and Social Communication 26.4.1 Maternal Behavior 26.4.2 Alloparental Care 26.4.3 Communication and Recognition 26.5 Population Regulation and Limits to Recovery 26.5.1 Predation, Neonatal Mortality and Disease 26.5.2 Fisheries and Human Interactions Including Marine Plastic Entanglement 26.6 Evolution of a Unique Reproductive Cycle 26.6.1 Breeding Cycle 26.6.2 Breeding Asynchrony and Nutrient Poor Environment 26.6.3 The Family Farm and Matrilineal Specialization 26.6.4 Prime Number Theory 26.6.5 Decoupling Embryonic Diapause from Photoperiod and Reducing Costs of Reproduction 26.7 The Future of Australian Sea Lions References Chapter 27: Australian Fur Seal: Adapting to Coexist in a Shared Ecosystem 27.1 Establishment and Distribution 27.2 Population Genetics 27.3 Description 27.4 Life-History 27.4.1 Territory Establishment and Defence 27.4.2 Gestation, Parturition and Breeding 27.4.3 Maternal Attendance and Lactation 27.4.4 Pup Growth 27.4.5 Pup Transition to Independence 27.5 Foraging Ecology 27.5.1 Ranges 27.5.2 Foraging Trip Strategies 27.5.3 Diving Behavior 27.5.4 Diet 27.6 Conservation 27.7 Conclusion References Chapter 28: Guadalupe Fur Seals and California Sea Lions: Two Sympatric Otariids from the California Current Ecosystem 28.1 Introduction 28.2 The Use of Stable Isotopes in Trophic Ecology 28.3 How Was This Work Done? 28.4 Our Findings 28.5 Final Remarks References Chapter 29: Fur Seal Trophic Ecology in Uruguay 29.1 South American Fur Seals in Uruguay 29.2 Diet 29.3 Individuals Prefer a Broad Diet 29.4 Females Travel Far and a Long Time; Pups Wait and Fast 29.5 Short, Shallow and Nocturnal Divers 29.6 Comparative Remarks References Chapter 30: Galapagos Sea Lions and Fur Seals, Adapted to a Variable World 30.1 Introduction 30.2 Galapagos Fur Seals 30.2.1 Foraging Ecology 30.2.2 Demographics 30.3 Galapagos Sea Lion 30.3.1 Species Description and Reproduction 30.3.2 Foraging Behavior and Feeding 30.3.3 Population Trends 30.4 Adaptations of Galapagos Otariids to Ecosystem Uncertainties References Index