Table of Contents

Part I. WHAT ARE AMPHIBIANS AND REPTILES?

Chapter 1. Why Study Herpetology?
1.1. Changing Perspectives
1.2. The Diversity of Amphibians and Reptiles
Amphibians
Reptiles
1.3. Shared Characteristics of Amphibians and Reptiles
Ectothermal thermoregulation
Costs and benefits of ectothermy and endothermy
Body size and shape
Ectothermy and efficiency
1.4. Amphibians and Reptiles in Terrestrial Ecosystems
1.5. The Future of Amphibians and Reptiles

Chapter 2. Phylogenetic Systematics and the Origins of Amphibians and Reptiles
2.1. Principles of Phylogenetics and Taxonomy
Building phylogenies
Rank-free taxonomy and phylogenetic nomenclature
Discovering and describing new species
Molecular data and species identification
2.2. Evolutionary Origins and Processes of Amphibian and Reptile Diversity
The ecological transition from water to land
The transition from fish to tetrapods
Early tetrapodomorphs
Early tetrapods
2.3. Three Hypotheses for the Origin of Extant Amphibians
The temnospondyl hypothesis
The lepospondyl hypothesis
The diphyly hypothesis
Why do different analyses support different hypotheses of lissamphibian origins?
2.4. Relationships among Extant Lissamphibian Lineages
Monophyly of Lissamphibia
Paedomorphosis in lissamphibian evolution
2.5. Characteristics and Origin of the Amniotes
The origins of Amniota
The major amniote lineages: Synapsida and Diapsida
2.6. Diapsida: Lepidosauria and Archosauria
Lepidosauria
Archosauria
2.7. The Debated Origins of Turtles

Chapter 3. Systematics and Diversity of Extant Amphibians
3.1. What Is an Amphibian?
Amphibian life histories
Amphibian skin
3.2. Caudata: Salamanders
Morphology
Reproduction and life history
Fossil record
Systematics and Phylogeny of Salamanders
3.3. Anura: Frogs and Toads
Skeletal morphology
Reproduction and life history
Fossil record
Systematics and Phylogeny of Frogs
3.4. Gymnophiona: Caecilians
Morphology
Reproduction and life history
Fossil record
Systematics and Phylogeny of Caecilians

Chapter 4. Systematics and Diversity of Extant Reptiles
4.1. Characteristics of Reptiles
Reptile skin
Sensory systems
4.2. Lepidosauria: Rhynchocephalia
The tuatara
Fossil record
4.3. Lepidosauria: Squamata
General anatomy of squamates
Reproduction and sex determination
Tail autotomy
Limb reduction
Venom and venom-delivering structures
Squamate phylogeny
Fossil record
4.4. Squamata: Lizards
Systematics and Phylogeny of Lizards
4.5. Squamata: Serpentes, the Snakes
Unique morphological features of snakes
Reproduction
Dentition
Fossil record
Systematics and Phylogeny of Snakes
4.6. Crocodylia
Fossil record
Systematics and Phylogeny of Crocodylians
4.7. Testudines: Turtles
The turtle skeleton
The turtle shell
Locomotion and reproduction
Fossil record
Systematics and Phylogeny of Turtles

Chapter 5. The Biogeography of Amphibians and Reptiles
5.1. Biogeographic Analysis
5.2. Dispersal
Transoceanic dispersal
Transcontinental dispersal
Human-mediated dispersal
5.3. Vicariance
Pangaea and the Mesozoic origin of modern amphibians and reptiles
Laurasian and Gondwanan origins of extant amphibians and reptiles
Africa and faunal exchange with southern Europe
Madagascar: An ancient continental fragment
The Seychelles Islands: Endemism on a microarchipelago
The Indian subcontinent: Gondwanan elements rafting to southern Asia
The Australian Plate: Mixed Gondwanan and Laurasian elements
South America
Origin of Central American assemblages
Merging faunas: The Great American Biotic Interchange
The Indo-Australian Archipelago and the Philippines
5.4. Phylogeography: Biogeography of the Recent Past
Climate change, isolation, and refugia
Sky islands
5.5. Island Biogeography
Island gigantism and dwarfism
Adaptive radiations on islands
Island paleoendemism

Part II. HOW DO THEY WORK?

Chapter 6. Water and Temperature Relations
6.1. Water Uptake and Loss
Routes of water gain
Routes of water loss
6.2. Water and the Ecology of Amphibians and Reptiles
Short-term water balance
Long-term water balance
6.3. Heat Gain and Loss
Absorption of solar radiation
Metabolic heat production, M
Infrared (thermal) radiative exchange, R
Convective exchange of heat, C
Evaporative cooling, LE
Conduction, G
6.4. Mechanisms of Thermoregulation
Heliothermy
Thigmothermy and kleptothermy
Set-point temperatures
6.5. Physiological Mechanisms of Thermoregulation
Moving heat within the body
Metabolic heat production
6.6. Effectiveness of Thermoregulation
6.7. The Coevolution of Energy and Water Exchange

Chapter 7. Energetics and Performance
7.1. Sites of Gas Exchange
Nonpulmonary gas exchange
Pulmonary gas exchange
Gas exchange by eggs
7.2. Patterns of Blood Flow
Pulmonary and systemic blood flow
Blood flow in the heart
Cardiac shunts
7.3. ATP Synthesis: Oxidative and Glycolytic Metabolism
Red and white muscle
Metabolic rates
Glycolytic metabolism
Lactic acid metabolism
Total ATP production and activity
7.4. Environmental Variables and Performance
Effects of environmental conditions on adults
Effects of the nest environment
7.5. Energy Costs of Natural Activities
Locomotion
Feeding
Vocalization by anurans
7.6. Metabolic Depression: Aestivation, Hibernation, and Freezing
Hibernation and aestivation
Freeze resistance and freeze tolerance
7.7. Annual Energy Budgets

Chapter 8. Reproduction and Life Histories of Amphibians
8.1. Sex Determination
8.2. Reproductive Cycles
8.3. Modes of Fertilization
Internal fertilization by salamanders
External fertilization by salamanders
External fertilization by anurans
Internal fertilization by anurans
Internal fertilization by caecilians
8.4. Hybridogenesis and Kleptogenesis
The European waterfrog complex
Kleptogenesis among Ambystoma
8.5. Reproductive Modes
Caecilians
Salamanders
Anurans
8.6. Evolution of Direct Development and Viviparity
8.7. Evolution of Parental Care
Benefits of parental care
Costs of parental care
Who cares?
8.8. Egg Size and Clutch Size
Variability in egg size
8.9. Complex Life Cycles, Larval Development, and Metamorphosis
Complex life cycles
Larval development of caecilians
Larval development of salamanders
Larval development of anurans
Hormonal control of metamorphosis and developmental plasticity
The ecology of metamorphosis
8.10. Paedomorphosis

Chapter 9. Reproduction and Life Histories of Reptiles
9.1. Sex Determination
Mechanisms of sex determination
Adaptive significance of GSD and TSD
Ecological consequences of TSD
9.2. Asexual Reproduction
9.3. Reproductive Modes
Oviparity: Eggshells, eggs, and nests
Characteristics of viviparous species
The evolution of viviparity
9.4. Parental Care
9.5. Reproductive Anatomy, Gametes, and Sperm Storage
9.6. Embryonic Development
Differentiation and growth
Genetic regulation of development
Phenotypic plasticity in development
Comparative developmental biology
9.7. Reproductive Cycles
Aseasonal cycles
Seasonal cycles
9.8. Life Histories of Reptiles
Patterns of life history variation
Trade-offs and life history evolution

Chapter 10. Body Support and Locomotion
10.1. Body Support and Thrust
10.2. Lever Systems
10.3. Terrestrial Locomotion with Limbs
Salamanders
Lizards
Crocodylians
Turtles
10.4. Jumping
10.5. Terrestrial Limbless Locomotion
10.6. Aquatic Locomotion
Undulatory swimmers
Oscillatory swimmers
10.7. Burrowing
10.8. Climbing
Grasping
Adhesion
10.9. Aerial Locomotion

Chapter 11. Feeding
11.1. Suction and Suspension Feeding
Salamanders and caecilians
Tadpoles
11.2. Terrestrial Feeding Mechanisms
Akinetic, nonprojectile feeding
Projectile feeding
Kinetic feeding
11.3. Capturing and Subduing Prey
Constriction
Envenomation
Digestion

Part III. WHAT DO THEY DO?

Chapter 12. Spatial Ecology
12.1. Ecological Consequences of Movement
12.2. Methods for Studying Movements
12.3. Types of Movement
12.4. Resource Dispersion and the Use of Space
Spatial strategies
Home range fidelity and homing
Finding the way home
12.5. Territoriality
Costs and benefits of territoriality
Sex and territoriality
Site defense
12.6. Migration
Breeding migrations of amphibians
Breeding migrations of terrestrial reptiles
Breeding migrations of sea turtles
Migrations to overwintering sites
12.7. Dispersal Strategies
Dispersal by amphibians
Dispersal by reptiles

Chapter 13. Communication
13.1. Modes of Communication
13.2. Constraints on Signal Production
Body size
Physiological constraints
Predation
13.3. Communication and Noise
Acoustic noise
Visual noise
13.4. Communication by Salamanders
Communication by plethodontids
Communication by salamandrids
13.5. Communication by Anurans
Acoustic communication
Visual communication
Chemical communication
13.6. Communication by Turtles
Tactile communication
Visual communication
Chemical communication
Acoustic communication
13.7. Communication by Crocodylians
Communication by alligators
Communication by crocodiles
Communication by juvenile crocodylians
Do crocodylians use chemical signals?
13.8. Communication by Squamates
Visual communication
Chemical communication
Acoustic communication

Chapter 14. Mating Systems and Sexual Selection
14.1. The Relationship of Mating Systems to Sexual Selection
14.2. Mating Systems of Amphibians and Reptiles
Scramble competition mating systems
Mate searching
Mate guarding
Multiple mate-guarding strategies
Leks and choruses
Resource defense
14.3. Variables Affecting Male Reproductive Success
Male persistence and allocation of resources
Male competitive ability
Female choice
Alternative mating tactics
Polyandry and sperm competition
14.4. Sexual Size Dimorphism

Chapter 15. Diets, Foraging, and Interactions with Parasites and Predators
15.1. Diets
Carnivory
Cannibalism
Herbivory and omnivory
Ontogenetic and sexual variation in diet
Temporal and spatial variation in diet
15.2. Amphibians and Reptiles as Consumers
Innate and learned responses to prey
Sensory modalities
Foraging modes
Foraging modes that don't fit the paradigm
Phylogeny and foraging modes
15.3. Parasites
Internal parasites
External parasites
15.4. Predators
Predation on eggs
Predation on amphibian larvae
Predation on postnatal amphibians and reptiles
15.5. Defensive Mechanisms
Avoiding detection
Signaling inedibility
Avoiding capture
Preventing consumption
15.6. Coevolution of Predators and Prey

Chapter 16. Populations and Species Assemblages
16.1. Population Ecology
What is a population?
Population parameters and population dynamics
Population change and population regulation
16.2. Community Ecology
Determinants of community structure
Patterns and mechanisms of amphibian and reptile assemblages
16.3. The Dynamic Nature of Assemblages
Impact of climate change
Recovery of assemblages following habitat destruction
16.4. Gradients in Species Richness
Latitudinal gradients
Elevational gradients

Part IV. WHAT ARE THEIR PROSPECTS FOR SURVIVAL?

Chapter 17. Conservation and the Future of Amphibians and Reptiles
17.1. Declining Populations of Amphibians and Reptiles
Amphibians
Reptiles
17.2. Major Themes in Biodiversity Conservation
17.3. Human Perceptions of Amphibians and Reptiles
17.4. Impact of Humans on Amphibians and Reptiles
Habitat modification and destruction
Introduction of exotic species
Pollution
Commercial exploitation for food
Commercial exploitation for skins, art, souvenirs, and other uses
Hallucinogens, hunting magic, and medicine
Pets
Research and teaching
Global climate change
Interaction among factors
17.5. Patterns of Species Extinction and Extirpation
Long-lived species
Species with low reproductive rates


Species that have poor dispersal and colonization abilities
Continental endemics
Oceanic island endemics
Species with colonial nesting habits
Migratory species
17.6. Conservation Options
Habitat protection
Coexistence with humans
Research
Education
National legislation
Control of international trade
Reestablishing populations
Farming and ranching
Sustainable harvesting
17.7. Declining Amphibians: A Model Issue
Possible causes of declines
Focus: Chytrid fungi and amphibian decline
17.8. Rediscovery and De-Extinction
Rediscovery of species
De-Extinction

Glossary
Literature Cited
Taxonomic Index
Subject Index

About the Author

F. Harvey Pough is Professor Emeritus in the Thomas H. Gosnell School of Life Sciences of the Rochester Institute of Technology. He received his Ph.D. in 1968 from the University of California at Los Angeles, with Kenneth S. Norris and Malcolm S. Gordon. In addition to Herpetology, he has headed up the author team on nine editions of Vertebrate Life (Benjamin Cummings/Pearson). Dr. Pough is a fellow of both the American Association for the
Advancement of Science and The Herpetologists' League, and Past President of the American Society of Ichthyologists and Herpetologists. His research focuses on organismal biology and evolutionary physiology, especially that of
amphibians and reptiles.

Robin M. Andrews is Professor Emerita in the Department of Biological Sciences at Virginia Tech. She received her Ph.D. in 1971 at the University of Kansas with Charles Michener and Daniel Janzen. She made the transition from Entomology to Herpetology during a postdoctoral fellowship at the Smithsonian Tropical Research Institute with A. Stanley Rand and Ernest Williams. Her current research interests are the physiological ecology and natural history of reptilian eggs and
embryos and the evolution and adaptive significance of developmental patterns of squamate reptiles.

Martha L. Crump is a behavioral ecologist who works with tropical amphibians in the areas of reproduction, ecology, and conservation. She is currently Adjunct Professor in the Department of Biology and the Ecology Center at Utah State University, and Adjunct Professor in the Department of Biology at Northern Arizona University. Dr. Crump received her Ph.D. from the University of Kansas in 1974, working with William E. Duellman. Her research, carried out in Amazonian Ecuador,
focused on community ecology and reproductive behaviors of frogs. In 1997, she received the Distinguished Herpetologist Award from The Herpetologists' League. Together with Dr. James P. Collins, Dr. Crump published
Extinction in Our Times: Global Amphibian Decline (2009).

Alan H. Savitzky is Professor and Head of the Department of Biology at Utah State University. He completed his graduate degrees at the University of Kansas (with William E. Duellman), receiving a Smithsonian Predoctoral Fellowship to conduct his dissertation research at the National Museum of Natural History. Dr. Savitzky is a Past President of both the Society for the Study of Amphibians and Reptiles and the American Society of Ichthyologists and Herpetologists, and currently
serves as Treasurer of the World Congress of Herpetology. His research concerns the integrative biology of amphibians and reptiles, especially snakes. Specific interests include the evolutionary morphology of feeding
and defensive structures, evolutionary development of sensory organs and glands, and, most recently, the evolution of chemical defenses in snakes.

Kentwood D. Wells is Professor of Ecology and Evolutionary Biology at the University of Connecticut. He received his Ph.D. in 1976 from Cornell University, with F. Harvey Pough. His book, The Ecology and Behavior of Amphibians (2007) was Best Single-Volume Science Reference Book for 2007 (Association of American Publishers Professional and Scholarly Publishing Division) and an Outstanding Academic Title for 2008 (Choice magazine). His 1977 paper on The social behaviour
of anuran amphibians (Animal Behaviour 25:666-693) was the first of 12 papers designated as most influential in the first 60 years of the journal. Dr. Wells researches the social behavior and communication of amphibians.

Matthew C. Brandley is an Australian Research Council DECRA postdoctoral fellow at the School of Biological Sciences, University of Sydney, Australia. He received his PhD in 2008 from the University of California, Berkeley. Dr. Brandley studies the phylogenetics and morphological evolution of vertebrate animals, especially lizards and snakes. He is particularly interested in how complex structures and unique body plans convergently evolve, and he studies these phenomena using a
combination of genomic, gene expression, anatomical, and phylogenetic tools. He lives in New South Wales Australia with his wife, son, two cats, and two axolotls.

Reviews

"This book includes a nearly complete overhaul of the systematics sections (including new distribution maps) to reflect numerous recent revisions. Where taxonomic incongruence or controversy exists, recent alternative hypotheses are reported, emphasizing the complexities of phylogenetic inference and persistent knowledge gaps rather than misleading the uninitiated reader with single, arbitrarily resolved trees. Those familiar with previous editions will be
immediately taken by the artistry of the images, both the extensive use of color photographs and the masterfully redrawn illustrations and data figures, which capture many fascinating aspects of the lives
of our favorite animals. Instructors, and especially the current generation of herpetology students, will welcome the online supplemental materials, which include links to websites with herpetological information, videos, and news stories relevant to each chapter."
--Robert E. Espinoza, California State University, Northridge
"I see this book as an invaluable reference and perhaps the most important feature of this revision is the authors' commitment to keeping the book fresh and up to date. . . . the improvements to an already outstanding reference were clearly worth the wait. Every herpetologist should own a well-worn copy of this volume." --Jimmy A. McGuire, The Quarterly Review of Biology