Health and Welfare of Captive Reptiles

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