Herpetology

723 Pages·2015·320.046 MB·English

by  F. Harvey Pough, Robin M. Andrews, Martha L. Crump, Alan H. Savitzky, Kentwood D. Wells, Matthew C. Brandley

Herpetology
■ FOURTH EDITION
*
■ • • * * w
I F. HARVEY POUGH
ROBIN M. ANDREWS • MARTHA L. CRUMP
ALAN H. SAVITZKY • KENTWOOD D. WELLS
MATTHEW C. BRANDLEY
F. Harvey Pough • Rochester Institute of Technology, emeritus
Robin M. Andrews • Virginia Tech, emerita
Martha L. Crump • Utah State University
Alan H. Savitzky • Utah State University
Kentwood D. Wells • University of Connecticut
Matthew C. Brandley • University of Sydney
Sinauer Associates, Inc. •
Publishers
Sunderland, Massachusetts • USA
About the Cover
A Neotropical reticulated glass frog (Hyalinobatrachium valerioi, Centrolenidae) guards a
clutch of eggs on the underside of a leaf overhanging a stream. Male glass frogs remain with
their clutches until they hatch, protecting them from egg-eating predators and releasing water
to keep the eggs hydrated. The frog's pattern of yellow spots surrounded by green resembles
the egg mass and may conceal it from predators such as birds. Photograph © Ingo Arndt/
Minden Pictures.
Herpetology, Fourth Edition
Copyright © 2016 by Sinauer Associates, Inc. All Rights Reserved. For information address
Sinauer Associates, Inc., PO Box 407, Sunderland, MA 01375 U.S.A.
FAX [phone]
orders@sinaue 匚 com
publish@sinaue 匚 com
www.sinaue 匚 com
Table of Contents and Part Opener Photographs:
p. vii: Alpine newt (Mesotriton alpestris), © Dirk Ercken/Shutterstock
p. ix: Shed snake skin, © Tyler Fox/Shutterstock
p. x: Reinwardt's gliding frog (Rhacophorus reinwardtil), © reptiles4all/Shutterstock
p. xii: Collared lizard (Crotaphytus bicinctores), © reptiles4all/Shutterstock
Part 1 top: Vietnamese blue flying frog (Rhacophorus helenae), © davemhuntphotography/
Shutterstock
Part 1 bottom: Western ringneck snake (Diadophis punctatus), © SuperStock/Alamy
Part 2 top: Lake Titicaca water frog (Jelmatobius culeus), © Pete Oxford/Nature Picture Library/
Corbis
Part 2 bottom: Green basilisk (Basiliscus plumifrons) running on water surface, © Bence Mate/
Nature Picture Library/Corbis
Part 3 top: Frilled lizard (Chlamydosaurus kingii) displaying, © Dave Watts/Alamy
Part 3 bottom: Moor frogs (Rana arvalis) in amplexus, © ImageBroker/Alamy
Part 4 top: Green tree python (Morelia viridis), © Siegfried Kuttig/ImageBroker/Corbis
Part 4 bottom: Green sea turtles (Chelonia mydas) and tourists, © Jarno Gonzalez
Zarraonandia/Shutterstock
Library of Congress Cataloging-in-Publication-Data
Pough, F. Harvey.
Herpetology / F. Harvey Pough, Rochester Institute of Technology, emeritus, Robin M.
Andrews, Virginiarlech, emerita, Martha L. Crump, Utah State University, Alan H. Savitzky,
Utah State University, Kentwood D. Wells, University of Connecticut, Matthew C. Brandley,
University of Sydney. -- Fourth edition.
pages cm
Includes bibliographical references and index.
ISBN 978-1-60535-233-6 (casebound)
1.Herpetology. I. Title.
QL641.H47 2015
597.9-dc23 2015017774
Printed in U.S.A.
6 5 4 3 2 1
Brief Contents
PART I ■ What Are Amphibians and Reptiles?
■ Why Study Herpetology? 3
CHAPTER 1
■ Phylogenetic Systematics and the Origins of Amphibians and Reptiles 19
CHAPTER 2
■ Systematics and Diversity of Extant Amphibians 41
CHAPTER 3
■ Systematics and Diversity of Extant Reptiles 107
CHAPTER 4
■ The Biogeography of Amphibians and Reptiles 203
CHAPTER 5
PART II ■ How Do They Work?
■ Water and Temperature Relations 227
CHAPTER 6
■ Energetics and Performance 259
CHAPTER 7
■ Reproduction and Life Histories of Amphibians 287
CHAPTER 8
Reproduction and Life Histories of Reptiles 321
CHAPTER 9 ■
Body Support and Locomotion 353
CHAPTER 10 ■
■ Leeding 379
CHAPTER 11
PART III ■ What Do They Do?
12 ■ Spatial Ecology 409
CHAPTER
■ Communication 437
CHAPTER 13
■ Mating Systems and Sexual Selection 475
CHAPTER 14
■ Diets, Eoraging, and Interactions with Parasites and Predators 501
CHAPTER 15
Populations and Species Assemblages 533
CHAPTER 16 ■
PART IV ■ What Are Their Prospects for Survival?
■ Conservation and the Future of Amphibians and Reptiles 557
CHAPTER 17
Contents
PART I ■ What Are Amphibians and Reptiles?
CHAPTER 1 The transition from fish to tetrapods 26
Why Study Herpetology? 3 Early tetrapodomorphs 28
Early tetrapods 28
1.1 Changing Perspectives 3 2.3 Three Hypotheses for the Origin of Extant
1.2 The Diversity of Amphibians and Reptiles 4 Amphibians 29
The temnospondyl hypothesis 29
Amphibians 4
The lepospondyl hypothesis 29
Reptiles 7
1.3 Shared Characteristics of Amphibians and The diphyly hypothesis 30
Reptiles 12 Why do different analyses support different hypotheses of
lissamphibian origins? 31
Ectothermal thermoregulation 13
2.4 Relationships among Extant Lissamphibian
Costs and benefits of ectothermy and endothermy 13
Lineages 32
Body size and shape 13
Monophyly of Lissamphibia 32
Ectothermy and efficiency 14
Paedomorphosis in lissamphibian evolution 33
1.4 Amphibians and Reptiles in Terrestrial
2.5 Characteristics and Origin of the Amniotes 35
Ecosystems 15
The origins of Amniota 35
1.5 The Future of Amphibians and Reptiles 16
The major amniote lineages: Synapsida and Diapsida 35
2.6 Diapsida: Lepidosauria and Archosauria 37
CHAPTER 2
Lepidosauria 37
Phylogenetic Systematics and the
Archosauria 38
Origins of Amphibians and Reptiles 19 2.7 The Debated Origins of Turtles 38
2.1 Principles of Phylogenetics and Taxonomy 19
CHAPTER 3
Building phylogenies 22
Systematics and Diversity of Extant
Rank-free taxonomy and phylogenetic nomenclature 23
Discovering and describing new species 24 Amphibians 41
Molecular data and species identification 24
3.1 What Is an Amphibian? 41
2.2 Evolutionary Origins and Processes of Amphibian
and Reptile Diversity 25 Amphibian life histories 41
The ecological transition from water to land 25 Amphibian skin 42
Contents vii
Reproduction 152
Dentition 153
Fossil record 153
Systematics and Phylogeny of Snakes 154
4.6 Crocodylia 180
Fossil record 181
Systematics and Phylogeny of Crocodylians 181
4.7 Testudines: Turtles 184
The turtle skeleton 185
3.2 Caudata: Salamanders 44 The turtle shell185
Locomotion and reproduction 186
Morphology 44
Fossil record 186
Reproduction and life history 44
Systematics and Phylogeny of Turtles 187
Fossil record 45
Systematics and Phylogeny of Salamanders 45
3.3 Anura: Frogs and Toads 55 CHAPTER 5
Skeletal morphology 55 The Biogeography of Amphibians
Reproduction and life history 56
and Reptiles 203
Fossil record 57
Systematics and Phylogeny of Frogs 59
5.1 Biogeographic Analysis 203
3.4 Gymnophiona: Caecilians 95
5.2 Dispersal 204
Morphology 95
Transoceanic dispersal 204
Reproduction and life history 97
Transcontinental dispersal 205
Fossil record 99
Human-mediated dispersal 205
Systematics and Phylogeny of Caecilians 99
5.3 Vicariance 206
Pangaea and the Mesozoic origin of modern amphibians
CHAPTER 4 and reptiles 207
Laurasian and Gondwanan origins of extant amphibians
Systematics and Diversity of Extant
and reptiles 208
Reptiles 107
Africa and faunal exchange with southern Europe 209
Madagascar: An ancient continental fragment 209
4.1 Characteristics of Reptiles 107
The Seychelles Islands: Endemism on a
Reptile skin 107 microarchipelago 210
Sensory systems 110 The Indian subcontinent: Gondwanan elements rafting to
4.2 Lepidosauria: Rhynchocephalia 114 southern Asia 210
The tuatara 114 The Australian Plate: Mixed Gondwanan and Laurasian
elements 210
Fossil record 115
South America 211
4.3 Lepidosauria: Squamata 115
Origin of Central American assemblages 212
General anatomy of squamates 116
Merging faunas: The Great American Biotic Interchange 212
Reproduction and sex determination 116
The Indo-Australian Archipelago and the Philippines 213
Tail autotomy 117
5.4 Phylogeography: Biogeography of the Recent
Limb reduction 118
Past 216
Venom and venom-delivering structures 119
Climate change, isolation, and refugia 216
Squamate phylogeny 119
Sky islands 218
Fossil record 120
5.5 Island Biogeography 219
4.4 Squamata: Lizards 122
Island gigantism and dwarfism 219
Systematics and Phylogeny of Lizards 122
Adaptive radiations on islands 220
4.5 Squamata: Serpentes, the Snakes 151
Island paleoendemism 221
Unique morphological features of snakes 151
viii Contents
PART II ■ How Do They Work?
CHAPTER 6 Lactic acid metabolism 273
Water and Temperature Relations 227 Total ATP production and activity 273
7.4 Environmental Variables and Performanee 274
6.1 Water Uptake and Loss 227 Effects of environmental conditions on adults 274
Routes of water gain 228 Effects of the nest environment 275
Routes of water loss 231 7.5 Energy Costs of Natural Activities 276
6.2 Water and the Ecology of Amphibians and Locomotion 277
Reptiles 236
Feeding 277
Short-term water balance 236 Vocalization by anurans 279
Long-term water balance 237 7.6 Metabolic Depression: Aestivation, Hibernation,
6.3 Heat Gain and Loss 239 and Freezing 281
Absorption of solar radiation, Qabs 240 Hibernation and aestivation 281
Metabolic heat production, M 242 Freeze resistance and freeze tolerance 281
Infrared (thermal) radiative exchange, R 242 7.7 Annual Energy Budgets 283
Convective exchange of heat, C 243
Evaporative cooling, LE 243
CHAPTER 8
Conduction, G 244
Reproduction and Life Histories of
6.4 Mechanisms of Thermoregulation 245
Amphibians 287
Heliothermy 245
Thigmothermy and kleptothermy 247
8.1 Sex Determination 287
Set-point temperatures 248
8.2 Reproductive Cycles 288
6.5 Physiological Mechanisms of Thermoregulation 250
8.3 Modes of Fertilization 289
Moving heat within the body 250
Internal fertilization by salamanders 289
Metabolic heat production 251
6.6 Effectiveness of Thermoregulation 252 External fertilization by salamanders 291
External fertilization by anurans 291
6.7 The Coevolution of Energy and Water
Exchange 254 Internal fertilization by anurans 293
Internal fertilization by caecilians 294
8.4 Hybridogenesis and Kleptogenesis 294
CHAPTER 7
The European waterfrog complex 294
Energetics and Performance 259
Kleptogenesis among Ambystoma 295
8.5 Reproductive Modes 296
7.1 Sites of Gas Exchange 259
Caecilians 296
Nonpulmonary gas exchange 259
Salamanders 296
Pulmonary gas exchange 261
Anurans 299
Gas exchange by eggs 265
8.6 Evolution of Direct Development and
7.2 Patterns of Blood Flow 266
Viviparity 305
Pulmonary and systemic blood flow 266
8.7 Evolution of Parental Care 306
Blood flow in the heart 266
Benefits of parental care 306
Cardiac shunts 269
Costs of parental care 307
7.3 ATP Synthesis: Oxidative and Glycolytic
Who cares? 307
Metabolism 270
8.8 Egg Size and Clutch Size 307
Red and white muscle 271
Variability in egg size 308
Metabolic rates 271
Glycolytic metabolism 272
Contents ix
&9 Complex Life Cycles, Larval Development, and Crocodylians 357
Metamorphosis 309 Turtles 357
Complex life cycles 309 10.4 Jumping 358
Larval development of caecilians 309 10.5 Terrestrial Limbless Locomotion 360
Larval development of salamanders 309 10.6 Aquatic Locomotion 363
Larval development of anurans 309
Undulatory swimmers 365
Hormonal control of metamorphosis and developmental
Oscillatory swimmers 365
plasticity 311
10.7 Burrowing 366
The ecology of metamorphosis 311
10.8 Climbing 369
8.10 Paedomorphosis 315
Grasping 369
Adhesion 370
CHAPTER 9 10.9 Aerial Locomotion 373
Reproduction and Life Histories of
Reptiles 321
CHAPTER 11
Feeding 379
9.1 Sex Determination 321
Mechanisms of sex determination 322
11.1 Suction and Suspension Feeding 379
Adaptive significance of GSD and TSD 323
Salamanders and caecilians 379
Ecological consequences of TSD 325
Tadpoles 381
9.2 Asexual Reproduction 327
11.2 Terrestrial Feeding Mechanisms 384
9.3 Reproductive Modes 328
Akinetic, nonprojectile feeding 384
Oviparity: Eggshells, eggs, and nests 328
Projectile feeding 387
Characteristics of viviparous species 330
Kinetic feeding 393
The evolution of viviparity 331
11.3 Capturing and Subduing Prey 398
9.4 Parental Care 332
Constriction 398
9.5 Reproductive Anatomy, Gametes, and Sperm
Envenomation 399
Storage 335
Digestion 403
9.6 Embryonic Development 337
Differentiation and growth 337
Genetic regulation of development 340
Phenotypic plasticity in development 340
Comparative developmental biology 341
9.7 Reproductive Cycles 342
Aseasonal cycles 343
Seasonal cycles 343
9.8 Life Histories of Reptiles 345
Patterns of life history variation 345
Trade-offs and life history evolution 346
CHAPTER 10
Body Support and Locomotion 353
10.1 Body Support and Thrust 353
10.2 Lever Systems 354
10.3 Terrestrial Locomotion with Limbs 356
Salamanders 356
Lizards 356
X Contents
PARTHI ■ What Do They Do?
CHAPTER 12 Visual communication 458
Spatial Ecology 409 Chemical communication 458
Acoustic communication 459
12.1 Ecological Consequences of Movement 409 13.7 Communication by Crocodylians 459
12.2 Methods for Studying Movements 410 Communication by alligators 459
12.3 Types of Movement 411 Communication by crocodiles 461
12.4 Resource Dispersion and the Use of Space 412 Communication by juvenile crocodylians 461
Do crocodylians use chemical signals? 461
Spatial strategies 412
13.8 Communication by Squamates 461
Home range fidelity and homing 418
Finding the way home 421 Visual communication 461
12.5 Territoriality 423 Chemical communication 468
Acoustic communication 471
Costs and benefits of territoriality 423
Sex and territoriality 423
Site defense 424 CHAPTER 14
12.6 Migration 425
Mating Systems and Sexual
Breeding migrations of amphibians 425
Selection 475
Breeding migrations of terrestrial reptiles 426
Breeding migrations of sea turtles 426
14.1 The Relationship of Mating Systems to Sexual
Migrations to overwintering sites 428 Selection 476
12.7 Dispersal Strategies 428
14.2 Mating Systems of Amphibians and Reptiles 477
Dispersal by amphibians 429
Scramble competition mating systems 478
Dispersal by reptiles 430
Mate searching 481
Mate guarding 482
CHAPTER 13 Multiple mate-guarding strategies 484
Communication 437 Leks and choruses 485
Resource defense 486
13.1 Modes of Communication 437 14.3 Variables Affecting Male Reproductive
Success 488
13.2 Constraints on Signal Production 438
Male persistence and allocation of resources 489
Body size 438
Male competitive ability 490
Physiological constraints 439
Female choice 492
Predation 439
Alternative mating tactics 493
13.3 Communication and Noise 440
Acoustic noise 440
Visual noise 441
13.4 Communication by Salamanders 442
Communication by plethodontids 442
Communication by salamandrids 444
13.5 Communication byAnurans 447
Acoustic communication 447
Visual communication 452
Chemical communication 456
13.6 Communication by Turtles 456
Tactile communication 457
Contents xi
Polyandry and sperm competition 496 15.5 Defensive Mechanisms 520
14.4 Sexual Size Dimorphism 498 Avoiding detection 520
Signaling inedibility 522
Avoiding capture 526
CHAPTER 15
Preventing consumption 527
Diets, Foraging, and Interactions with
15.6 Coevolution of Predators and Prey 529
Parasites and Predators 501
CHAPTER 16
15.1 Diets 501
Carnivory 501 Populations and Species
Cannibalism 503 Assemblages 533
Herbivory and omnivory 505
Ontogenetic and sexual variation in diet 506 16.1 Population Ecology 533
Temporal and spatial variation in diet 507 What is a population? 533
15.2 Amphibians and Reptiles as Consumers 508 Population parameters and population dynamics 534
Innate and learned responses to prey 508 Population change and population regulation 537
Sensory modalities 509 16.2 Community Ecology 538
Foraging modes 511 Determinants of community structure 538
Foraging modes that don't fit the paradigm 513 Patterns and mechanisms of amphibian and reptile
Phylogeny and foraging modes 514 assemblages 539
16.3 The Dynamic Nature of Assemblages 546
15.3 Parasites 515
Internal parasites 515 Impact of climate change 546
Recovery of assemblages following habitat destruction 547
External parasites 516
16.4 Gradients in Species Richness 548
15.4 Predators 518
Predation on eggs 518 Latitudinal gradients 549
Predation on amphibian larvae 519 Elevational gradients 551
Predation on postnatal amphibians and reptiles 519
PART IV ■ What Are Their Prospects for
Survival?
CHAPTER 17 Introduction of exotic species 564
Conservation and the Future of Pollution 566
Commercial exploitation for food 568
Amphibians and Reptiles 557
Commercial exploitation for skins, art, souvenirs, and other
uses 568
17.1 Declining Populations of Amphibians and
Hallucinogens, hunting magic, and medicine 569
Reptiles 557
Pets 570
Amphibians 557
Research and teaching 571
Reptiles 559
Global climate change 571
17.2 Major Themes in Biodiversity Conservation 559
Interaction among factors 572
17.3 Human Perceptions of Amphibia ns and
17.5 Patterns of Species Extinction and Extirpation 573
Reptiles 560
Long-lived species 573
17.4 Impact of Humans on Amphibia ns and
Reptiles 561 Species with low reproductive rates 574
Species that have poor dispersal and colonization
Habitat modification and destruction 562
abilities 574