Health and Welfare of Captive Reptiles (Chapman and Hall Materials Management/)

This extensively revised and expanded new edition offers concepts, principles and applied information that relates to the wellbeing of reptiles. As a manual on health and welfare in a similar vein to volumes addressing the sciences of anatomy, behaviour or psychology, this book thoroughly examines the biology of reptile welfare and is about meeting biological needs. The editors, acknowledged experts in their own right, have once again drawn together an extremely impressive international group of contributors. Positive and negative implications of general husbandry and research programs are discussed. In addition to greatly revised original content are nine new chapters offering readers novel insight into: • sensory systems • social behaviour • brain and cognition • controlled deprivation and enrichment • effects of captivity-imposed noise and light disturbance on welfare • spatial and thermal factors• evidential thresholds for species suitability in captivity • record keeping as an aid to captive care • arbitrary husbandry practices and misconceptions The authors have adopted a user-friendly writing style to accommodate a broad readership. Although primarily aimed at academic professionals, this comprehensive volume is fundamentally a biology book that will also inform all involved in captive reptile husbandry. Among others, zoo personnel, herpetologists, veterinarians, lab animal scientists, and expert readers in animal welfare and behavioural studies will benefit from this updated work. Acknowledgements Contents Editors and Contributors 1: Introduction 1.1 Authors 1.2 Peer Review and Quality Control 1.3 Audience 1.4 Introducing Chapters 1.5 Conclusion 2: Physiology and Functional Anatomy 2.1 Introduction 2.2 Body Temperature, Energetics, and Ectothermy 2.2.1 Temperature and Energy Expenditure 2.2.2 Regulation of Body Temperature 2.2.3 Variation of Body Temperature 2.2.4 Functional Significance of Body Temperature Variation 2.3 Light and Photoreception 2.4 Water Exchange and Humidity 2.4.1 Water Exchange 2.4.2 Water Loss and Integument 2.5 Ecdysis 2.6 Osmoregulation and Nitrogen Metabolism 2.7 Digestive Physiology and Nutrition 2.8 Respiration and Circulation 2.8.1 Lung Structure and Function 2.8.2 Blood Circulation 2.9 Pain and Stress 2.10 Animal Welfare Considerations References 3: Sensory Systems 3.1 Introduction 3.2 Photoreception 3.2.1 Eyes and Vision 3.2.2 Visual Communication 3.2.3 Non-Visual Photoreception 3.2.3.1 The Parietal Eye and Circadian Rhythms 3.2.3.2 Skin and Photoreception 3.3 Mechanoreception 3.3.1 Cutaneous Mechanoreceptors and Touch 3.3.1.1 Tactile Communication 3.3.1.2 Hydrodynamic Sense 3.4 Ears and Hearing 3.4.1 Acoustic Communication 3.4.2 Vestibular System and Equilibrium 3.5 Chemoreception 3.5.1 Tongue and Gustation 3.5.2 Nose and Olfaction 3.5.3 Vomeronasal System 3.5.4 Chemical Communication 3.6 Thermoreception 3.6.1 Heat-Sensing Pits in Snakes 3.7 Magnetoreception 3.8 Animal Welfare Considerations References 4: Biology of Stress 4.1 Introduction 4.2 Defining Stressors, Stress, and the Stress Response 4.2.1 Stress Response Pathways 4.2.1.1 The HPA Axis 4.2.1.2 The SAM Axis 4.2.2 Colour Change as a Stress Indicator 4.2.3 Stress and Reproduction 4.2.4 Stress and Immunity 4.2.5 Example Studies of Stress Response in Reptiles 4.3 Future Directions 4.4 Animal Welfare Conclusions Appendix: Abbreviations References 5: Normal Behaviour 5.1 Introduction 5.2 Maintenance Behaviours 5.2.1 Maintenance: Feeding Behaviour 5.2.2 Feeding Behaviour: Animal Welfare Conclusions 5.2.3 Maintenance: Thermoregulatory Behaviour 5.2.4 Thermoregulatory Behaviour: Animal Welfare Conclusions 5.3 Distance-Reducing Behaviour 5.3.1 Distance-Reducing Behaviour: Aggregation 5.3.2 Aggregation: Animal Welfare Conclusions 5.3.3 Distance-Reducing Behaviour: Courtship and Mating Behaviour 5.3.4 Courtship and Mating Behaviour: Animal Welfare Conclusions 5.3.5 Distance-Reducing Behaviour: Parental Care 5.3.6 Parental Care: Animal Welfare Conclusions 5.4 Agonistic Behaviour 5.4.1 Agonistic Behaviour: Territoriality and Combat 5.4.2 Territoriality and Combat: Animal Welfare Conclusions 5.4.3 Agonistic Behaviour: Antipredator Behaviour 5.4.4 Antipredator Behaviour: Animal Welfare Conclusions 5.5 Animal Welfare Conclusions References 6: Social Behaviour as a Challenge for Welfare 6.1 Introduction 6.2 Brief Taxonomic Breakdown of Social Behaviour in Reptiles 6.3 The Welfare of Captive Reptiles and Its Assessment 6.3.1 Social Factors and Welfare 6.4 Problems and Future Directions 6.5 Animal Welfare Conclusions References 7: Brains, Behaviour, and Cognition: Multiple Misconceptions 7.1 Introduction 7.2 Misconception 1: Reptile Brains Are Ten Times Smaller than Those of Birds and Mammals of Similar Body Size 7.3 Misconception 2: Reptile Brains Lack a Cerebral Cortex 7.4 Misconception 3: Reptilian Behaviour Is Simple, Stereotyped, Uninteresting, and Primitive 7.5 Misconception 4: Reptiles Are Cognitive Underachievers 7.6 Misconception 5: Reptiles Are Solitary, Lack Parental Care, and Do Not Form Stable Social Relationships 7.7 Misconception 6: Reptiles Do Not Play 7.8 Misconception 7: Reptiles Do Not Dream 7.9 Further Directions on the Study of the Reptile Brain and Cognition 7.10 Animal Welfare Implications References 8: Psychological and Behavioural Principles and Problems 8.1 Introduction 8.2 Living Wild and Noticing Captivity 8.3 Domestication and Non-Domestication 8.4 Maladaptation to Captivity 8.5 Stereotyped Behaviours and Maladaptive Stereotypies 8.6 Recognising and Interpreting Signs of Psychological and Ethological Well-being and Poor Welfare 8.7 Ontogenetic Processes and Innate (Unlearnt) Traits in Nature 8.7.1 Anticipating Ontogenetic Processes and Traits in the Captive Environment 8.8 Psychological, Ethological, Physiological, and Physical Interrelationships 8.9 Stress and Stressors 8.9.1 Behavioural Indicators of Stress 8.9.1.1 Signs of Positive and Negative Psychological and Behavioural States 8.10 Consciousness and Self-Awareness 8.10.1 Awareness of Self-Preservation and Well-Being 8.11 Specific Psychological and Ethological Problems 8.11.1 Exploratory, Search, and Escape Activities 8.11.2 Interaction with Transparent Boundaries 8.11.3 Hyperactivity 8.11.4 Hypoactivity 8.11.5 Restricted by Provision 8.11.6 Social Stress 8.11.7 Aggression 8.11.8 Disposition-Related Voluntary Hypothermia 8.11.9 Feeding Behaviour and Problems 8.11.9.1 Excessive Weight Gain and Loss 8.11.9.2 Co-Occupant Harassment and Competitive and Non-Competitive Feeders 8.11.9.3 Abnormal Ingestion of Items 8.11.9.4 Post-Feeding Quiescence 8.11.9.5 Cannibalism 8.12 General Considerations 8.12.1 Establishing Possible Meanings from Behavioural Signs 8.12.2 Body Posture and Position 8.12.3 Biological and Behavioural Strategies in Health and Disease 8.12.4 Similarities and Differences Between Juveniles and Adults 8.12.5 Reproduction 8.12.6 Disturbance of Rest and Sleep 8.13 Literature Developments 8.14 Animal Welfare Conclusions References 9: Effects of Ontogeny, Rearing Conditions, and Individual Differences on Behaviour: Welfare, Conservation, and Invasive Speci... 9.1 Introduction 9.2 The Embryonic Period 9.3 Parental Care 9.4 Enclosure Temperature 9.5 Handling, Enclosures, and Novel Environments 9.6 Cage Structure 9.7 Social Arrangements 9.8 Feeding 9.9 Antipredator Behaviour 9.10 Individuality, Plasticity, and Long-Term Influence of Captive Regimes 9.11 Animal Welfare Conclusions References 10: Controlled Deprivation and Enrichment 10.1 Introduction 10.2 Controlled Deprivation 10.3 Enrichment 10.4 Enrichment as Applied to Captive Reptiles 10.5 Enrichment or Basic Husbandry?: A Philosophical Quandary 10.6 A Model for Reptile Enrichment 10.7 Examples of Reptile Enrichment 10.7.1 Environmental Enrichment 10.7.1.1 Thermal Regimes 10.7.1.2 Water Relations 10.7.1.3 Substrates 10.7.1.4 Enclosure Furnishings 10.7.1.5 Refugia 10.7.1.6 Spatial Considerations 10.7.2 Sensory Enrichment 10.7.2.1 Chemoreception 10.7.2.2 Visual Considerations 10.7.2.3 Tactile Considerations 10.7.2.4 Acoustic Considerations 10.7.3 Dietary Enrichment 10.7.4 Cognitive Enrichment 10.7.4.1 Novel Objects 10.7.4.2 Problem-Solving Tasks 10.7.4.3 Training 10.7.5 Social Enrichment 10.8 Evaluating Enrichment 10.9 Challenges Associated with Reptile Enrichment 10.10 Animal Welfare Conclusions References 11: Effects of Noise and Light 11.1 Introduction 11.1.1 Basic Characteristics of Sound and Noise Perception 11.1.1.1 Hearing Ranges and Effects of Anthropogenic Noise on Different Orders 11.1.1.2 Characteristics of Noise in Captivity and Possible Noise Control Measures 11.1.1.3 Sound and Vibrations 11.1.2 Basic Characteristics of Light Perception: Special Characteristics 11.1.2.1 Light Perception and Effects of Light Disturbance on Various Orders 11.1.2.2 Characteristics of Light in Captivity and Possible Light Control Measures 11.2 Animal Welfare Conclusions References 12: Ethologically Informed Design and DEEP Ethology in Theory and Practice 12.1 Introduction 12.2 Ethological Attitude 12.3 Deep Ethology 12.3.1 Development 12.3.2 Ecology 12.3.3 Evolution 12.3.4 Physiology 12.4 Ethologically Informed Practice 12.5 Validity 12.6 Laboratory and Field 12.7 Observation and Context 12.8 Animal Models 12.9 Case Studies 12.9.1 Thermoregulation 12.9.2 Stress 12.9.3 Social Dominance 12.9.4 Reproduction 12.10 Mitigation of Distress 12.11 Animal Welfare Conclusions References 13: Spatial and Thermal Factors 13.1 Introduction 13.2 Historical Approaches 13.3 Space Utilisation, Exploration, and Enrichment 13.3.1 Home Ranges 13.3.2 Rack Systems 13.3.3 Space Utilisation: Sociality and Size 13.3.4 Overcrowding 13.3.5 How Much Space Is Enough Space? 13.4 Thermal Considerations 13.4.1 Thermal Gradients and Zones 13.4.2 Subtle Thermal Changes May Be Critical 13.5 Animal Welfare Conclusions References 14: Nutritional Considerations 14.1 Introduction 14.2 Selection of Food 14.3 Acquisition of Food 14.4 Water Supply 14.5 Processing, Assimilation, and Elimination 14.6 Target Nutrient Values for Appropriate Diets 14.7 Considerations for Provision of UV Light 14.8 Miscellaneous Factors and their Effects on Nutrition 14.9 Future Research Opportunities 14.10 Animal Welfare Conclusions References 15: Naturalistic Versus Unnaturalistic Environments 15.1 Introduction 15.2 Terminology for Types of Environment 15.3 Naturalistic Versus Unnaturalistic 15.4 Naturalistic Environments 15.4.1 General Captive Conditions 15.4.2 Research Conditions 15.4.3 Summary Conclusion 15.5 Unnaturalistic Environments 15.5.1 Veterinary Management 15.5.2 Quarantine 15.5.3 Summary Conclusion 15.6 Practical Considerations 15.6.1 Access to Animals 15.6.2 Maintenance Efficiency and Hygiene 15.7 Animal Welfare Conclusions References 16: Evidential Thresholds for Species Suitability in Captivity 16.1 Introduction 16.2 Information, Compliance, and Knowledge Deficits 16.3 Essential Principles for Animal Welfare 16.4 Assessing the Suitability of Reptiles for Captivity 16.5 Suitability Tools 16.5.1 Example Existing Suitability Models 16.5.2 Proposed Suitability Tools 16.5.2.1 Using the Algorithm-Based Tools: Difficulties and Weaknesses of the Self-Assessment Approach 16.5.2.2 Instructions for Use of the Algorithm-Based Tools 16.5.2.3 Worked Examples 16.6 Conclusions 16.7 Animal Welfare Conclusions Appendix: Worked Examples of the EMODE Method for Table 16.2 References 17: Record Keeping as an Aid to Captive Care 17.1 Introduction 17.2 Historical Records 17.3 Identification Records 17.4 Husbandry Records 17.4.1 Reproductive Data 17.4.2 Behavioural Data 17.5 Veterinary and Pathology Records 17.6 Record Keeping Systems 17.7 Record Keeping for Retrospective and Comparative Studies 17.8 Animal Welfare Conclusions References 18: Arbitrary Husbandry Practices and Misconceptions 18.1 Introduction 18.2 Evidential Assessment 18.3 General Misconceptions (See Table 18.1) 18.3.1 `Reptiles are Easy to Keep ́ 18.3.2 `Reptiles Live Longer in Captivity than in the Wild ́ 18.3.3 `Reptiles Would Not Feed, Grow and Reproduce Unless Experiencing Good Welfare ́ 18.3.4 `Reptiles are Only Active When Seeking Food or a Mate, and If These Features are Provided, Nothing Else is Required ́ 18.4 Environmental Considerations (See Table 18.2) 18.4.1 `UV is Unnecessary for Many Commonly Kept Reptiles ́ 18.4.2 `Arbitrary or Single Thermal Conditions are Acceptable as Long as Some Heat Source is Provided: All Heat is the Same ́ 18.4.3 `Adequate Thermal Gradients are Possible in Small Enclosures ́ 18.4.4 `Single, Constant Humidity Levels are Acceptable ́ 18.5 Nutritional Considerations (See Table 18.3) 18.5.1 `Food is Food Whatever its Origins ́ 18.5.2 `Arbitrary Feeding Quantities and Frequencies are Acceptable ́ 18.5.3 `Standing Water Should Not be Provided for Xeric Habitat Species ́ 18.5.4 `Arbitrary Placement of Water Sources is Acceptable ́ 18.6 Behavioural Considerations (See Table 18.4) 18.6.1 `Reptiles are Not Social ́ 18.6.2 `Snakes Thrive in Small Enclosures ́ 18.6.3 `Reptiles (Especially Snakes) are Sedentary and Do Not Use Space ́ 18.6.4 `Snakes Suffer from Insecurity, Agoraphobia or Anorexia in Large Environments ́ 18.6.5 `Snakes Do Not Need to Stretch Out ́ 18.7 Animal Welfare Conclusions References 19: Miscellaneous Factors 19.1 Introduction 19.2 Stress, Pain and Sensitivity 19.2.1 Stress 19.2.2 Natural Versus Captivity-Related Stressors and Stress 19.2.3 Stressors and Stress in Nature and in Captivity 19.2.4 Stress and Natural Rhythms 19.2.5 Stress Measurement Through Disease and Mortality Data 19.2.6 Emotional Stress 19.3 Pain Perception and Assessment Sensitivity 19.3.1 Stress-Induced Analgesia 19.4 Thermal Factors, Thermoregulation and Light 19.4.1 Captive Reptiles and the Thermal Environment 19.4.2 Thermal Burns 19.4.3 Light and Photo-Invasive Environments 19.5 Circadian Rhythms and Nocturnalism 19.6 Growth 19.7 Electromagnetism in the Artificial Environment 19.8 Reintroductions to Nature 19.9 Euthanasia and Killing 19.9.1 Problems in Establishing Signs of Life and Death 19.9.2 Euthanasia and Killing Methods 19.9.3 Decapitation and Spinal Cord Severance 19.9.4 Hypothermia in Relation to Anaesthesia, Euthanasia and Killing 19.10 Occupancy and Post-Occupancy Evaluation 19.11 Human-Animal Interactions and Relationships 19.11.1 Changing Human Behaviour to Improve Captive Welfare 19.12 Ethical Considerations 19.13 Animal Welfare Conclusions References Index Animal Index Introduction: health and welfare of captive reptiles / Clifford Warwick, Fredric L. Frye, and James B. Murphy Physiology and functional anatomy / Harvey B. Lillywhite and Robert E. Gatten, Jr. Biology of stress: interactions with reproduction, immunology and intermediary metabolism / Louis J. Guilette, Jr., Alison Cree and Andrew A. Rooney Nutritional considerations / Fredric L. Frye Veterinary perspectives and techniques in husbandry and research / John E. Cooper and David L. Williams Naturalistic versus clinical environments in husbandry and research / Clifford Warwick and Catrina Steedman Normal behaviour / James C. Gillingham Effects of ontogenetic processes and rearing conditions / Gordon M. Burghardt and Donna Layne Behavioural consequences of husbandry manipulations: indicators of arousal, quiescence, and environmental awareness / David Chiszar ... [et al.] Psychological and behavioural principles and problems / Clifford Warwick Ethologically informed design in husbandry and research / Neil Greenberg Miscellaneous factors affecting health and welfare / Phillip C. Arena and Clifford Warwick This most important book fully examines the welfare of captive reptiles and discusses the positive and negative implications of general husbandry and research programmes. The editors, acknowledged experts in their own right, have drawn together an extremely impressive international group of contributors providing clearly written and comprehensive accounts of aspects such as physiology, physical stress, diet, veterinary and environmental issues, normal behaviour, psychological stress and informed design in research.