Vertebrate Life

729 Pages·2013·37.885 MB·English

by  F. Harvey Pough, Christine M. Janis, John B. Heiser

F. HARVEY POUGH
Rochester Institute of Technology
CHRISTINE M. JANIS
Brown University
JOHN B. HEISER
Cornell University
VERTEBRATE
LIFE
NINTH EDITION
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ISBN 10: 0-321-77336-5
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About the Authors
F. Harvey Pough is a Professor of Biology at the Rochester Institute of Technology.
He began his biological career at the age of fourteen when he and his sister studied
the growth and movements of a population of eastern painted turtles in Rhode
Island. His research now focuses on organismal biology, blending physiology,
morphology, behavior, and ecology in an evolutionary perspective. Undergraduate
students regularly participate in his research and are coauthors of many of his pub-
lications. He especially enjoys teaching undergraduates and has taught introductory
biology and courses in vertebrate zoology, functional ecology, herpetology, environ-
mental physiology, and animal behavior. He has published more than a hundred
papers reporting the results of field and laboratory studies of turtles, snakes, liz-
ards, frogs, and tuatara that have taken him to Australia, New Zealand, Fiji, Mexico,
Costa Rica, Panama, and the Caribbean, as well as most parts of the United States.
He is a Fellow of the American Association for the Advancement of Science and a
past President of the American Society of Ichthyologists and Herpetologists.
Christine M. Janis is a Professor of Biology at Brown University, where she
teaches comparative anatomy and vertebrate evolution and is a recipient of the
Elizabeth Leduc Prize for Distinguished Teaching in the Life Sciences. A British
citizen, she obtained her bachelor's degree at Cambridge University and then
crossed the pond to get her Ph.D. at Harvard University. She is a vertebrate pale-
ontologist with a particular interest in mammalian evolution and faunal respons-
es to climatic change. She first became interested in vertebrate evolution after
seeing the movie Fantasia at the impressionable age of seven. That critical year
was also the year that she began riding lessons, and she has owned at least one
horse since the age of twelve. Many years later, she is now an expert on ungulate
(hoofed mammal) evolution and has recently expanded her interests to the evolu-
tion of the Australian mammal fauna, especially the kangaroos. She is a Fellow of
the Paleontological Society and is currently President of the Society for the Study
of Mammalian Evolution. She attributes her life history to the fact that she has
failed to outgrow either the dinosaur phase or the horse phase.
John B. Heiser was born and raised in Indiana and completed his undergraduate
degree in biology at Purdue University. He earned his Ph.D. in ichthyology from
Cornell University for studies of the behavior, evolution, and ecology of coral reef
fishes, research that he continues today with colleagues specializing in molecular
biology. For fifteen years, he was Director of the Shoals Marine Laboratory operated
by Cornell University and the University of New Hampshire on the Isles of Shoals
in the Gulf of Maine. While at the Isles of Shoals, his research interests focused on
opposite ends of the vertebrate spectrum—hagfish and baleen whales. He enjoys
teaching vertebrate morphology, evolution, and ecology, both in the campus class-
room and in the field, and is a recipient of the Clark Distinguished Teaching Award
from Cornell University. His hobbies are natural history, travel and nature pho-
tography, and videography, especially underwater using scuba. He has pursued his
natural history interests on every continent and all the world's major ocean regions.
Because of his experience, he is a popular ecotourism leader, having led Cornell
Adult University groups to the Caribbean, Sea of Cortez, French Polynesia, Central
America, the Amazon, Borneo, Antarctica, and Spitsbergen in the High Arctic.
iii
Brief Contents
Vertebrate Diversity, Function, and Evolution 1
PART I
1 The Diversity, Classification, and Evolution of Vertebrates 2
2 Vertebrate Relationships and Basic Structure 19
3 Early Vertebrates: Jawless Vertebrates and the Origin of Jawed Vertebrates 47
Non-Amniotic Vertebrates: Fishes and Amphibians 71
PART II
4 Living in Water 72
5 Radiation of the Chondrichthyes 99
6 Dominating Life in Water: The Major Radiation of Fishes 122
7 Geography and Ecology of the Paleozoic Era 158
8 Living on Land 167
9 Origin and Radiation of Tetrapods 189
10 Salamanders, Anurans, and Caecilians 211
Sauropsida: Turtles, Lepidosaurs, and Archosaurs 253
PART III
11 Synapsids and Sauropsids: Two Approaches to Terrestrial Life 254
12 Turtles 287
13 The Lepidosaurs: Tuatara, Lizards, and Snakes 310
14 Ectothermy: A Low-Cost Approach to Life 349
15 Geography and Ecology of the Mesozoic Era 364
16 Mesozoic Diapsids: Dinosaurs, Crocodilians, Birds, and Others 371
17 Avian Specializations 407
Synapsida: The Mammals 447
PART IV
18 The Synapsida and the Evolution of Mammals 448
19 Geography and Ecology of the Cenozoic Era 471
20 Mammalian Diversity and Characteristics 480
21 Mammalian Specializations 513
22 Endothermy: A High-Energy Approach to Life 537
23 Body Size, Ecology, and Sociality of Mammals 559
24 Primate Evolution and the Emergence of Humans 581
25 The Impact of Humans on Other Species of Vertebrates 614
Appendix A-1
Glossary G-1
Credits C-1
Index I-1
iv
Contents
Preface xv
Vertebrate Diversity, Function, and Evolution 1
PART I
1

The Diversity, Classification, and Evolution of Vertebrates 2
1.1 The Vertebrate Story 2
1.2 Classification of Vertebrates 8
1.3 Phylogenetic Systematics 8
1.4 The Problem with Fossils: Crown and Stem Groups 11
1.5 Evolutionary Hypotheses 12
1.6 Earth History and Vertebrate Evolution 15
Summary 16
Discussion Questions 16
Additional Information 17
2
Vertebrate Relationships and Basic Structure 19
2.1 Vertebrates in Relation to Other Animals 19
2.2 Definition of a Vertebrate 24
2.3 Basic Vertebrate Structure 25
Summary 45
Discussion Questions 45
Additional Information 46
3
Early Vertebrates: Jawless Vertebrates and the Origin
of Jawed Vertebrates 47
3.1 Reconstructing the Biology of the Earliest Vertebrates 47
3.2 Extant Jawless Fishes 50
3.3 The Importance of Extant Jawless Vertebrates in Understanding
Ancient Vertebrates 56
3.4 The Radiation of Paleozoic Jawless Vertebrates—“Ostracoderms” 57
3.5 The Basic Gnathostome Body Plan 57
3.6 The Transition from Jawless to Jawed Vertebrates 62
3.7 Extinct Paleozoic Jawed Fishes 66
Summary 68
Discussion Questions 69
Additional Information 69
v
Non-Amniotic Vertebrates: Fishes and Amphibians 71
PART II
4
Living in Water 72
4.1 The Aquatic Environment 72
4.2 Water and the Sensory World of Fishes 77
4.3 The Internal Environment of Vertebrates 84
4.4 Exchange of Water and Ions 84
4.5 Responses to Temperature 90
4.6 Body Size and Surface/Volume Ratio 94
Summary 96
Discussion Questions 97
Additional Information 97
5
Radiation of the Chondrichthyes 99
5.1 Chondrichthyes—The Cartilaginous Fishes 99
5.2 Evolutionary Diversification of Chondrichthyes 102
5.3 The Paleozoic Chondrichthyan Radiation 102
5.4 The Early Mesozoic Elasmobranch Radiation 105
5.5 Extant Lineages of Elasmobranchs 106
5.6 Batoidea: Skates and Rays 116
5.7 Holocephali—The Little Known Chondrichthyans 118
Summary 119
Discussion Questions 119
Additional Information 121
6
Dominating Life in Water: The Major Radiation of Fishes 122
6.1 The Origin of Bony Fishes 122
6.2 Evolution of the Actinopterygii 126
6.3 Extant Actinopterygii—Ray-Finned Fishes 131
6.4 Locomotion in Water 137
6.5 Actinopterygian Reproduction 141
6.6 The Adaptable Fishes—Teleosts in Contrasting Environments 147
6.7 Conservation of Fishes 150
6.8 Sarcopterygii—The Lobe-Finned Fishes 153
Summary 156
Discussion Questions 156
Additional Information 157
7
Geography and Ecology of the Paleozoic Era 158
7.1 Earth History, Changing Environments, and Vertebrate Evolution 158
7.2 Continental Geography of the Paleozoic 159
7.3 Paleozoic Climates 162
7.4 Paleozoic Terrestrial Ecosystems 163
vi Contents
7.5 Paleozoic Extinctions 165
Additional Information 166
8
Living on Land 167
8.1 Support and Locomotion on Land 167
8.2 Eating on Land 175
8.3 Reproduction on Land 178
8.4 Breathing Air 178
8.5 Pumping Blood Uphill 178
8.6 Sensory Systems in Air 182
8.7 Conserving Water in a Dry Environment 184
8.8 Controlling Body Temperature in a Changing Environment 184
Summary 187
Discussion Questions 188
Additional Information 188
9
Origin and Radiation of Tetrapods 189
9.1 Tetrapod Origins 189
9.2 Radiation and Diversity of Non-Amniotic Paleozoic Tetrapods 197
9.3 Amniotes 201
Summary 208
Discussion Questions 209
Additional Information 209
10
Salamanders, Anurans, and Caecilians 211
10.1 Amphibians 211
10.2 Diversity of Life Histories of Amphibians 222
10.3 Amphibian Metamorphosis 237
10.4 Exchange of Water and Gases 238
10.5 Poison Glands and Other Defense Mechanisms 243
10.6 Mimicry 246
10.7 Why Are Amphibians Vanishing? 246
Summary 250
Discussion Questions 250
Additional Information 251
PART III Sauropsida: Turtles, Lepidosaurs, and Archosaurs 253
11
Synapsids and Sauropsids: Two Approaches to
Terrestrial Life 254
11.1 Taking Advantage of the Opportunity for Sustained
Locomotion 255
11.2 Increasing Gas Exchange: The Trachea and Lungs 259
Contents vii
11.3 Transporting Oxygen to the Muscles: Structure of the Heart 265
11.4 Taking Advantage of Wasted Energy: Endothermy 266
11.5 Getting Rid of Wastes: The Kidneys and Bladder 271
11.6 Sensing and Making Sense of the World: Eyes, Ears, Tongues, Noses, and Brains 281
Summary 284
Discussion Questions 284
Additional Information 285
12
Turtles 287
12.1 Everyone Recognizes a Turtle 287
12.2 But What Is a Turtle? Phylogenetic Relationships of Turtles 290
12.3 Turtle Structure and Function 291
12.4 Ecology and Behavior of Turtles 298
12.5 Reproductive Biology of Turtles 300
12.6 Hatching and the Behavior of Baby Turtles 302
12.7 Conservation of Turtles 306
Summary 307
Discussion Questions 308
Additional Information 308
13
The Lepidosaurs: Tuatara, Lizards, and Snakes 310
13.1 The Lepidosaurs 310
13.2 Radiation of Sphenodontids and the Biology of Tuatara 311
13.3 Radiation of Squamates 312
13.4 Ecology and Behavior of Squamates 321
13.5 Behavioral Control of Body Temperatures by Ectotherms 338
13.6 Temperature and Ecology of Squamates 342
Summary 346
Discussion Questions 347
Additional Information 347
14
Ectothermy: A Low-Cost Approach to Life 349
14.1 Vertebrates and Their Environments 349
14.2 Dealing with Dryness—Ectotherms in Deserts 350
14.3 Coping with Cold—Ectotherms in Subzero Conditions 356
14.4 The Role of Ectothermic Tetrapods in Terrestrial Ecosystems 359
Summary 362
Discussion Questions 362
Additional Information 363
15
Geography and Ecology of the Mesozoic Era 364
15.1 Mesozoic Continental Geography 365
15.2 Mesozoic Terrestrial Ecosystems 366
viii Contents
15.3 Mesozoic Climates 368
15.4 Mesozoic Extinctions 369
Additional Information 370
16
Mesozoic Diapsids: Dinosaurs, Crocodilians, Birds, and Others 371
16.1 The Mesozoic Fauna 371
16.2 Characteristics of Diapsids 372
16.3 Marine Lineages 372
16.4 Semiaquatic and Terrestrial Diapsids: Crocodylomorpha 379
16.5 The First Evolution of Flight: Pterosauria 382
16.6 Dinosaurs 384
16.7 Terrestrial Herbivores: Ornithischian and Sauropod Saurischian Dinosaurs 388
16.8 Terrestrial Carnivores: Theropod Dinosaurs 397
16.9 The Second Evolution of Flight: Birds 401
Summary 404
Discussion Questions 404
Additional Information 405
17
Avian Specializations 407
17.1 Early Birds and Extant Birds 407
17.2 The Structure of Birds 410
17.3 Wings and Flight 415
17.4 The Hindlimbs 422
17.5 Feeding and Digestion 425
17.6 Sensory Systems 429
17.7 Social Behavior 432
17.8 Mating Systems 434
17.9 Oviparity 435
17.10 Nests 437
17.11 Orientation and Navigation 440
17.12 Migration 441
Summary 444
Discussion Questions 445
Additional Information 445
Synapsida: The Mammals 447
PART IV
18
The Synapsida and the Evolution of Mammals 448
18.1 The Origin of Synapsids 448
18.2 Diversity of Nonmammalian Synapsids 450
18.3 Evolutionary Trends in Synapsids 455
Contents ix