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Vertebrate Life
TENTH EDITION
 Vertebrate
Life TENTH EDITION

F. Harvey Pough Christine M. Janis

Rochester Institute of Technology, Brown University, Emerita,
Emeritus and University of Bristol

Chapter 26 “Primate Evolution and the Emergence of Humans”

by Sergi López-Torres, University of Toronto Scarborough and
Roman Kozlowski Institute of Paleobiology, Polish Academy of Sciences

SINAUER ASSOCIATES
NEW YORK OXFORD
OXFORD UNIVERSITY PRESS

About the Cover
Artist’s reconstruction of two of the
earliest mammals, Morganucodon (left)
and Kuehneotherium (right). Both of
these Early Jurassic mammals were
insectivores about the size of extant
shrews. (Painting by John Sibbick,
2013, © Pamela Gill.)
Vertebrate Life, 10th Edition
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Library of Congress Cataloging-in-Publication Data
Names: Pough, F. Harvey, author. | Janis, Christine M. (Christine Marie),
1950- author.
Title: Vertebrate life / F. Harvey Pough, Rochester Institute of Technology,
Emeritus, Christine M. Janis, Brown University, Emerita, University of
Bristol ; chapter 26 “Primate Evolution and the Emergence of Humans” by
Sergi López-Torres, University of Toronto Scarborough, Roman Kozlowski
Institute of Paleobiology, Polish Academy of Sciences
Description: Tenth edition. | New York : Oxford University Press, [2019] |
Revised edition of: Vertebrate life / F. Harvey Pough, Christine M. Janis,
John B. Heiser. 9th ed. Boston : Pearson, c2013. | Includes
bibliographical references and index.
Identifiers: LCCN 2018005330 (print) | LCCN 2018005884 (ebook) | ISBN
9781605357218 (ebook) | ISBN 9781605356075 (casebound)
Subjects: LCSH: Vertebrates--Textbooks. | Vertebrates, Fossil--Textbooks.
Classification: LCC QL605 (ebook) | LCC QL605 .P68 2019 (print) | DDC
596--dc23
LC record available at https://lccn.loc.gov/2018005330
987654321
Printed in the United States of America
Brief Contents
CHAPTER 1 Evolution, Diversity, and Classification of Vertebrates 1
CHAPTER 2 What Is a Vertebrate? 19
CHAPTER 3 Jawless Vertebrates and the Origin of Jawed Vertebrates 41
CHAPTER 4 Living in Water 65
CHAPTER 5 Geography and Ecology of the Paleozoic Era 83
CHAPTER 6 Radiation and Diversification of Chondrichthyes 95
CHAPTER 7 Extant Chondrichthyans 103
CHAPTER 8 Radiation and Diversity of Osteichthyes 121
CHAPTER 9 Extant Bony Fishes 133
CHAPTER 10­ Origin and Radiation of Tetrapods 161
CHAPTER 11 Extant Amphibians 181
CHAPTER 12 Living on Land 211
CHAPTER 13 Geography and Ecology of the Mesozoic Era 233
CHAPTER 14 Synapsids and Sauropsids 241
CHAPTER 15 Ectothermy: A Low-Energy Approach to Life 269
CHAPTER 16 Turtles 283
CHAPTER 17 Lepidosaurs 301
CHAPTER 18 Crocodylians 329
CHAPTER 19 Mesozoic Diapsids: Dinosaurs and Others 343
CHAPTER 20 Endothermy: A High-Energy Approach to Life 371
CHAPTER 21 The Origin and Radiation of Birds 387
CHAPTER 22 Extant Birds 399
CHAPTER 23 Geography and Ecology of the Cenozoic Era 435
CHAPTER 24 Synapsida and the Evolution of Mammals 451
CHAPTER 25 Extant Mammals 481
CHAPTER 26 Primate Evolution and the Emergence of Humans 519
Contents
Preface xv
Chapter 1
Evolution, Diversity, and Classification
of Vertebrates 1
1.1 The Vertebrate Story 1
Major extant groups of vertebrates 2 The skeletomuscular system 30
Non-amniotes 2
Amniotes 4
1.2 Classification of Vertebrates 5
Binominal nomenclature 5
Phylogenetic systematics 5
Applying phylogenetic criteria 6
Morphology-based and molecular-based phylogenies
Using phylogenetic trees 8
1.3 Crown and Stem Groups 9 Jawless Vertebrates and the Origin
1.4 Genetic Mechanisms of Evolutionary Change 9
Phenotypes and fitness 10
Developmental regulatory genes 11
Intragenerational versus transgenerational phenotypic
modification 14
1.5 Earth History and Vertebrate Evolution 14
Chapter 2
What Is a Vertebrate? 19
2.1 Vertebrates in Relation to Other Animals 19
2.2 Characteristics of Chordates 20
Chordate origin and evolution 21
Extant nonvertebrate chordates 21
2.3 What Distinguishes a Vertebrate? 23
2.4 Vertebrate Embryonic Development 24
Development of the body 24
Development of the pharyngeal region 25
Development of the brain 27
Unique developmental features of vertebrates 27
2.5 Basic Vertebrate Structures 29
Adult tissue types 30
Mineralized tissues 30
2.6 Basic Vertebrate Systems 32
The alimentary system 32
The cardiovascular system 33
The excretory and reproductive systems 35
The sense organs 35
7
Chapter 3
of Jawed Vertebrates 41
3.1 The Earliest Evidence of Vertebrates 41
The origin of bone and other mineralized tissues 42
The mysterious conodonts 43
The environment of early vertebrate evolution 43
3.2 Cyclostomes: The Extant Jawless Fishes 46
Characters of cyclostomes 46
Fossil cyclostomes 46
Extant hagfishes: Myxiniformes 46
Lampreys: Petromyzontiformes 48
Cyclostomes and humans 49
3.3 Ostracoderms: Extinct Jawless Fishes 50
Characters of ostracoderms 50
Ostracoderm evolutionary patterns 50
   Contents  vii

3.4 The Basic Gnathostome Body Plan 51 Chapter 5

Gnathostome biology 51
What about soft tissues? 52 Geography and Ecology of the
3.5 The Origin of Jaws 52 Paleozoic Era 83
Early hypotheses of jaw origins 52 5.1 Shifting Continents and Changing Climates 83
The importance of the nose 55
5.2 Continental Geography of the Paleozoic 84
Developmental studies of extant vertebrates 55
Transitional anatomy in fossils 56 5.3 Paleozoic Climates 85
The selective value of jaws 56 5.4 Paleozoic Ecosystems 86
3.6 The Origin of Paired Appendages 56 Aquatic life 86
The advantages of fins 56 Terrestrial floral ecosystems 88
Fin development and the lateral somitic frontier 56 Terrestrial faunal ecosystems 89
Origin of the neck region 57 5.5 Paleozoic Extinctions 90
3.7 Extinct Paleozoic Jawed Fishes 58
Placoderms: Armored fishes 58 Chapter 6
Acanthodians 60
The surviving gnathostome groups 61
Radiation and Diversification of
Chondrichthyes 95
Chapter 4 6.1 Chondrichthyes: The Cartilaginous Fishes 95
Distinctive characters of chondrichthyans 95
Living in Water 65
6.2 
Evolutionary Diversification of
4.1 The Aquatic Environment 65 Chondrichthyes 98
Obtaining oxygen from water: Gills 65 Paleozoic chondrichthyan radiations 98
Obtaining oxygen from air: Lungs and other respiratory The Mesozoic chondrichthyan radiation 100
structures 66 Paleozoic and Mesozoic chondrichthyan
Adjusting buoyancy 67 paleobiology 101
4.2 Water and the Sensory World of Aquatic
Vertebrates 69 Chapter 7
Vision 69
Hearing 69
Extant Chondrichthyans 103
Chemosensation: Taste and smell 70 7.1 Morphology of Extant Chondrichthyans 103
Detecting water displacement 70 Skeleton 103
Electrical discharge 72 Jaws 103
Electroreception by sharks and rays 72 Skin 104
4.3 The Internal Environment of Vertebrates 74 7.2 Sharks (Selachii) 106
4.4 Exchange of Water and Ions 75 Sensory systems and prey detection 106
Nitrogen excretion 75 Ecology of sharks 108
The vertebrate kidney 75 Heterothermy 109
Regulation of ions and body fluids 76 Feeding 109
4.5 Vertebrates in Different Environments 76 Reproduction 110
Marine vertebrates 76 7.3 Skates and Rays (Batoidea) 112
Freshwater vertebrates: Teleosts and amphibians 78 Morphology 112
Euryhaline vertebrates 79 Ecology 112
Terrestrial vertebrates 80 Courtship and reproduction 113
7.4 Chimaeras (Holocephali) 113
7.5 Declining Shark Populations:
An Ecological Crisis 114
viii  Contents

Chapter 8
Radiation and Diversity of
Osteichthyes 121
8.1 The Origin of Bony Fishes 121
Earliest osteichthyans and the major groups
of bony fishes 123
8.2 Evolution of Actinopterygii 125
Basal actinopterygians 125
Neopterygii 125
Evolution of jaw protrusion 126
Pharyngeal jaws 127
Specializations of fins 128
8.3 Evolution of Sarcopterygii
Actinistia 129
Dipnoi 129
Tetrapodomorpha 130
Chapter 10
Origin and Radiation of Tetrapods 161
10.1 Tetrapod Origins 161
Tetrapodomorph fishes 161
Earliest tetrapods of the Late Devonian 163
10.2 Moving onto Land 165
Terrestrial and walking fishes today 165
How are fins made into limbs? 166
Body support and locomotion 167
Lung ventilation and dermal bone 168
10.3 Radiation and Diversity of Non-Amniote
Tetrapods 169
128 10.4 Amniotes 172
Derived features of amniotes 173
The amniotic egg 174
Patterns of amniote temporal fenestration 175

Chapter 9 Chapter 11
Extant Bony Fishes 133 Extant Amphibians 181
9.1 Actinopterygians: Ray-Finned Fishes 133 11.1 Diversity of Lissamphibians 181
Non-teleosts 133 Salamanders 181
Teleosts 134 Anurans 184
9.2 Swimming 139 Caecilians 188
Minimizing drag 141 11.2 Life Histories of Amphibians 188
Steering, stopping, and staying in place 141 Salamanders 189
9.3 Actinopterygian Reproduction 142 Anurans 191
Oviparity 142 The ecology of tadpoles 196
Viviparity 143 Caecilians 197
9.4 The Sex Lives of Teleosts 144 11.3 Amphibian Metamorphosis 198
Protandry 144 11.4 Exchange of Water and Gases 198
Protogyny 145 Cutaneous respiration 198
Hermaphroditism 146 Blood flow in larvae and adults 199
All-female species 147 Cutaneous permeability to water 199
9.5 Teleosts in Different Environments 147 Behavioral control of evaporative water loss 201
Deep-sea fishes 147 Uptake and storage of water 202
Coral reef fishes 152 11.5 Toxins, Venoms, and Other Defense
9.6 Heterothermal Fishes 152 Mechanisms 202
Warm muscles 152 Skin glands 202
Hot eyes 153 Toxicity and diet 203
Venomous amphibians 204
9.7 Sarcopterygians:Lobe-Finned Fishes 154
Actinistians: Coelacanths 154 11.6 Why Are Amphibians Vanishing? 205
Dipnoans: Lungfishes 155 Disease 205
Synergisms 206
9.8 Pollution, Overfishing, and Fish Farming 155
Freshwater fishes 155
Marine fishes 156
   Contents  ix

Chapter 12 14.2 Limb-Powered Locomotion 243

The basal amniote ankle joint 245
Living on Land 211 The sauropsid ankle joint 245
The synapsid ankle joint 245
12.1 Support and Locomotion on Land 211
The skeleton 211 14.3 Increasing Gas Exchange 247
The cranial skeleton 212 Synapsid lungs 248
The axial skeleton: Vertebrae and ribs 212 Sauropsid lungs 248
Axial muscles 213 The respiratory system of birds 248
The appendicular skeleton: limbs and limb girdles 215 Why are synapsid and sauropsid lungs so different? 248
Size and scaling 217 14.4 Transporting Oxygen to the Muscles:
Locomotion 217 The Heart 250
12.2 Eating on Land 219 14.5 The Evolution of Endothermy 253
How did endothermy evolve? 253
12.3 Breathing Air 220
Evaluating the models 254
12.4 Pumping Blood Uphill 220
When did endothermy evolve? 254
12.5 Sensory Systems in Air 222
14.6 Getting Rid of Wastes: The Kidneys 255
Vision 222
Nitrogen excretion by synapsids: The mammalian
Hearing 224 kidney 256
Olfaction 225 Nitrogen excretion by sauropsids: Renal and extrarenal
Proprioception 226 routes 260
12.6 Conserving Water in a Dry Environment 226 14.7 Sensing and Making Sense of the World 263
12.7 Controlling Body Temperature in a Changing Vision 263
Environment 227 Chemosensation: Gustation and olfaction 263
Ectothermy 227 Hearing 264
Endothermy 228 Brains 264
Ectothermy, endothermy, and heterothermy 228
Chapter 15
Chapter 13 Ectothermy: A Low-Energy Approach to
Geography and Ecology of the Life 269
Mesozoic Era 233
15.1 Vertebrates and Their Environments 269
13.1 Continental Geography of the Mesozoic 233 15.2 Dealing with Dryness: Ectotherms in
13.2 Mesozoic Climates 233 Deserts 269
Desert tortoises 270
13.3 Mesozoic Aquatic Life 234
The chuckwalla 271
13.4 Mesozoic Terrestrial Ecosystems 235 Desert amphibians 274
The Triassic 235
15.3 Coping with Cold: Ectotherms in Subzero
The Jurassic 236
Conditions 275
The Cretaceous 236
Frigid fishes 275
13.5 Mesozoic Extinctions 238 Frozen frogs 276
15.4 Energetics of Ectotherms and Endotherms 277
Chapter 14 Body size 277
Body shape 278
Synapsids and Sauropsids 241
Conversion efficiency 279
14.1 The Conflict between Locomotion and 15.5 The Role of Ectotherms in Terrestrial
Respiration 241 Ecosystems 279
Locomotion and lung ventilation of synapsids 242
Locomotion and lung ventilation of sauropsids 243
x  Contents

Chapter 16 17.8 Reproductive Modes 320

Sex determination 320
Turtles 283 Oviparity and viviparity 320
Parthenogenesis 321
16.1 Everyone Recognizes a Turtle 283
Parental care 321
Shell and skeleton 283
Families of extant turtles 285 17.9 Thermal Ecology 322
Organismal performance and temperature 322
16.2 Turtle Structure and Function 287
Lung ventilation 287 17.10 Lepidosaurs and Climate Change 323
The heart 288
Patterns of circulation and respiration 289 Chapter 18
Body size and temperature regulation 289
Crocodylians 329
16.3 Reproductive Biology of Turtles 289
Moisture and egg development 289 18.1 Diversity of Extant Crocodylians 329
Temperature-dependent sex determination 290 18.2 The Crocodylomorph Lineage 331
Parental care 291 Notosuchia 331
Hatching and the behavior of baby turtles 291 Neosuchia 332
16.4 Social Behavior, Communication, and 18.3 Predatory Behavior and Diet of Extant
Courtship 293 Crocodylians 333
16.5 Navigation and Migrations 293 18.4 Communication and Social Behavior 335
Navigation by adult turtles 294
18.5 Reproduction and Parental Care 336
Navigation by hatchling and juvenile sea turtles 295
Temperature-dependent sex determination 336
16.6 The Fateful Life-History Characteristics of Parental care 336
Turtles 295
18.6 The Skin Trade 337

Chapter 17
Chapter 19
Lepidosaurs 301
Mesozoic Diapsids: Dinosaurs and
17.1 Rhynchocephalians and the Biology Others 343
of Tuatara 301
19.1 Characteristics of Diapsids 343
17.2 Radiation of Squamates 303
Lizards 303 19.2 Diversity of Mesozoic Diapsids 343
Snakes 306 19.3 Lepidosauromorphs: Marine Diapsids 346
17.3 Foraging Modes 308 Terrestrial lepidosauromorphs 346
Correlates of foraging mode 309 Marine lepidosauromorphs 346
17.4 Skull Kinesis 312 19.4 Metriorhynchid Crocodylomorphs 349
17.5 Feeding Specializations of Snakes 312 19.5 Pterosaurs: The First Flying Vertebrates 349
Venom and fangs 314 The structure of pterosaurs 349
Hearts and stomachs 315 Reproduction, eggs, and parental care 352
Did the evolution of birds doom the pterosaurs? 352
17.6 Predator Avoidance and Defense 315
Crypsis 315 19.6 Triassic Faunal Turnover 352
“Eavesdropping” 316 19.7 The Structure and Function of Dinosaurs 353
Deterrence 316 Hips and legs 353
Autotomy 317 Dinosaur lineages 354
Venomous and poisonous snakes 317 19.8 Ornithischian Dinosaurs 355
17.7 Social Behavior 318 Thyreophora 355
Territoriality 318 Marginocephalia 357
Sociality 320 Ornithopoda 358
   Contents  xi

Social behavior of ornithischian dinosaurs 358 21.2 The Mosaic Evolution of Birds 392
Nesting and parental care by ornithischians 360 How—and why—birds got off the ground 392
19.9 Herbivorous Saurischians 360 The appearance of powered avian flight 393
The structure of sauropods 360 21.3 Early Birds 394
Social behavior of sauropods 362 21.4 The Mesozoic Radiations of Birds 394
Nesting and parental care by sauropods 362
19.10 Carnivorous Saurischians 363 Chapter 22
Tyrannosauroides 363
Ornithomimisauria 363 Extant Birds 399
Maniraptora 364
22.1 The Structure of Birds: Specialization
Social behavior of theropods 365 for Flight 399
Nesting and parental care by theropods 365 Feathers 401
19.11 Gigantothermy and the Body Temperature Streamlining and weight reduction 403
of Dinosaurs 366 Skeleton 403
Muscles 404
Chapter 20 22.2 Wings and Flight 404
Endothermy: A High-Energy Approach Wing muscles 406
Wing shape and flight characteristics 407
to Life 371
22.3 Feet 408
20.1 Balancing Heat Production with Heat Loss 371 Hopping, walking, and running 408
Whole-body metabolism 372 Climbing 409
Shivering and non-shivering thermogenesis 372 Swimming 409
Insulation 372
22.4 Feeding and Digestion 410
Evaporative cooling 373
Beaks, skulls, and tongues 410
20.2 Endotherms in the Cold 373 The digestive system 413
Avoiding cold and sharing heat 374
22.5 Sensory Systems 414
20.3 Facultative Hypothermia 375 Vision 415
Seasonal hypothermia 375 Hearing 416
Rest-phase hypothermia 375 Olfaction 416
Hibernation 376
22.6 Social Behavior 417
20.4 Endotherms in the Heat 378 Plumage colors and patterns 417
Temperature stress and scarcity of water 379 Vocalization, sonation, and visual displays 419
Strategies for desert survival 379
22.7 Oviparity 421
Avoidance 380
Egg biology 421
Relaxation of homeostasis by hyperthermia 381
Sex determination 422
Hypothermia in the desert 383
Maternal control of sex of offspring 422
22.8 Monogamy: Social and Genetic 423
Chapter 21
22.9 Nests and Parental Care 423
The Origin and Radiation of Birds 387 Incubation 423
Parental care 425
21.1 Avian Characters in Nonavian Theropods 387 Brood parasitism 425
Skeletal characters 387
22.10 Orientation and Navigation 426
Feathers 388
Reproduction and parental care 390 22.11 Migration 427
Body size 391 Migratory movements 427
Costs and benefits of migration 427
xii  Contents
22.12 Birds and Urbanization 428
Success in the city 429
Noise pollution 429
Not so sexy in the city 429
Chapter 23
Geography and Ecology of the
Cenozoic Era 435
23.1 Continental Geography of the Cenozoic
23.2 Cenozoic Climates 436
Paleogene and Neogene climates 436
The Pleistocene ice ages 438
23.3 Cenozoic Terrestrial Ecosystems 439
23.4 Biogeography of Cenozoic Mammals 441
The isolation of Australian mammals 441
The isolation of mammals on other continents 444
23.5 Cenozoic Extinctions 446
Chapter 24
Synapsida and the Evolution of
Mammals 451
24.1 The Origin of Synapsids 451
24.2 The Diversity of Non-Mammalian
Synapsids 451
Pelycosaurs: Basal non-mammalian synapsids 452
Therapsids: More derived non-mammalian
synapsids 456
Therapsid diversity 459
24.3 Evolutionary Trends in Synapsids 460
Evolution of the diaphragm 462
Evolution of a double occipital condyle 462
Evolution of jaws and ears 462
24.4 The First Mammals 465
Metabolic and growth rates 466
Skeletomuscular system 466
Feeding and mastication 467
Brain, senses, and behavior 468
The integument: Epidermis and glands 469
Lactation and suckling 471
Food processing and swallowing 472
Facial musculature 472
Internal anatomy 473
24.5 Mesozoic Mammals 474
Dual radiations of Mesozoic mammals 474
Chapter 25
Extant Mammals 481
25.1 Major Lineages of Mammals 481
Multituberculates 482
Monotremes 482
Marsupials 485
Placentals 487
25.2 Differences between Therians and Non-
435 Therians 491
Craniodental features 491
Postcranial skeletal features 492
Gait and locomotion 493
Hearing 494
Vision 494
Lactation 495
Information from the genes 495
Sex determination and sex chromosomes 496
25.3 Differences between Marsupials and
Placentals 496
25.4 Mammalian Reproduction 497
Mammalian urogenital tracts 498
Genitalia 498
Reproductive mode of monotremes: Matrotrophic
oviparity 499
Reproductive mode of therians: Matrotrophic
viviparity 500
The earliest therian condition, and the discredited notion
of placental superiority 502
25.5 Specializations for Feeding:
Teeth and Jaws 503
Mammalian teeth 503
Differences between carnivorous and herbivorous
mammals 503
Rodents: Specialized feeders 505
25.6 Specializations for Locomotion 506
Cursorial limb morphology 506
Fossorial limb morphology 507
25.7 The Evolution of Aquatic Mammals 508
Morphological adaptations for life in water 508
The evolution of cetaceans 509
25.8 Trophy Hunting 513
Endangering the endangered: The effect of perceived
rarity 514
The extinction vortex 515
   Contents  xiii

Chapter 26 What happened to the humans who were already

there? 543
Primate Evolution and the Emergence 26.5 Evolution of Human Characteristics 543
of Humans 519 Bipedalism 543
Large brains 544
26.1 Primate Origins and Diversification 519
Speech and language 545
Evolutionary trends and diversity in primates 519
Loss of body hair and development of skin
26.2 Origin and Evolution of Hominoidea 526 pigmentation 545
Diversity and social behavior of extant apes 527 Human technology and culture 546
Relationships within Hominoidea 531 26.6 Why Is Homo sapiens the Only Surviving
Diversity of fossil hominoids 531 Hominin Species? 546
26.3 Origin and Evolution of Humans 533 Hybridization among species of Homo 547
Early hominins 533 26.7 Humans and Other Vertebrates 547
Ecological and biogeographic aspects of early hominin Humans as superpredators and environmental
evolution 537 disruptors 547
26.4 Derived Hominins: The Genus Homo 537 Megafaunal extinctions 548
Homo erectus and Homo ergaster 537 Is this the Anthropocene? 549
The Dmanisi hominins 539
Homo floresiensis 539
Appendix A-1
Homo naledi 540 Glossary G-1
Precursors of Homo sapiens 540
The Neandertals 541 Illustration Credits IC-1
The Denisovans 542
Origins of modern humans 542 Index I-1
Preface
The sustainability of populations, and even the continued
existence of some species of vertebrates, is becoming ever
more problematic. In addition to overarching events that
affect all living organisms—such as global climate change
and acidification of the seas—each lineage of vertebrates
faces threats that are intimately entwined with the biolog-
ical characteristics of that lineage.
Thus, as we have prepared this tenth edition of Vertebrate
Life, the study of vertebrates has taken on new urgency.
Many areas of vertebrate biology have seen enormous ad-
vances since the previous edition:
• Phylogenies based on molecular data increasingly sup-
plement, reinforce, and in some cases contradict phy-
logenies based on morphology. We have incorporated
this information, including some of the cases where the
two approaches have generated different hypotheses
about the timing or sequence of evolutionary change.
• Our understanding of the genetic and epigenetic
control of development has advanced greatly, and this
evo-devo perspective provides a mechanistic under-
standing of evolutionary changes in phenotype.
• Newly discovered fossil sites, fossils, and even fossils
that preserve soft tissue structures have added enor-
mous detail (and often greater complexity) to our un-
derstanding of evolutionary lineages, especially early
tetrapods, feathered nonavian dinosaurs, and humans.
This edition reflects these changes. What has not changed
is the authors’ view of vertebrates as complex and fascinat-
ing organisms that can best be understood by considering
the interactions at multiple levels of biological organization
that shape the biology of a species. This book presents ver-
tebrates in a way that integrates all facets of their biology—
from anatomy and physiology to ecology and behavior—in
an evolutionary context
Authors
Two changes of authorship have occurred in this edition.
Ill health has compelled John B. Heiser to withdraw from
active participation. John has been an author since the first
edition of Vertebrate Life, enriching the chapters on fishes
with the experience gained from his field work in nearly
every ocean. We thank him for his contributions, and wish
him well.
We are delighted to have gained the assistance of Sergi
López-Torres who authored the chapter on primates and
the evolution of humans. Primate evolution is an extraor-
dinarily active field of research, and Sergi brings the depth
and breadth of knowledge of human evolution needed for
an overview of this increasingly complex field.
Organization
The scope of vertebrate biology and of evolutionary time
is vast, and encapsulating this multiplicity of themes and
mountains of data into a book or a semester course is a con-
tinuing challenge. Following suggestions from users, the
text has been extensively reorganized to improve the flow
of information. Topics have been split or merged, and some
have been deleted, resulting in presentation of information
in more manageable segments. Each chapter includes a list
of sources that will be useful to students, and all of the
sources that we consulted in preparing this edition, as well
as many sources from earlier editions, are available on the
book’s web page, oup.com/us/vertebratelife10e.
Sources
We have relied on these sources for the numbers of extant
species and their common and scientific names.
• FishBase: http://www.fishbase.se/search.php
• AmphibiaWeb: https://amphibiaweb.org/
• The Reptile Database: http://www.reptile-database.org/
• IOC World Birds list: http://www.worldbirdnames.org/
• Avibase: http://avibase.bsc-eoc.org/checklist.jsp
• Mammal Species of the World: https://www.depart-
ments.bucknell.edu/biology/resources/msw3/
• ASM Mammal Diversity Database: https://mammal-
diversity.org/
• IUCN Red List of threatened species: http://www.
iucnredlist.org/
xvi  Preface

Acknowledgments
Authors are only the visible tip of the iceberg that is a textbook. Expert librarians are
essential to any scholarly undertaking, and we are fortunate to have had the outstand-
ing assistance of Adwoa Boateng and Morna Hilderbrand (Wallace Library, Rochester
Institute of Technology) and Sue O’Dell (Hatch Science Library, Bowdoin College).
The figure legends cite the many artists and photographers who kindly allowed
us to use their work, often provided additional information, and in some cases took
new photos at our request.

We are grateful to the many colleagues who answered questions and suggested sources
for data and photographs:
Albert Bennett Margaret Fusari Stewart Nicol Wade C. Sherbrooke
Lucille Betti-Nash Harry W. Greene Martin Nyffler Rick Shine
Daniel Blackburn Katrina Halliday Kouki Okagawa Matthew Simon
Caleb M. Brown James Hanken Todd Pierson Megan Southern
Grover Brown Karsten E. Hartel Theodore Pietsch Zachary Stahlschmidt
Larry Buckley Gene Helfman Christopher Raxworthy Hyla Sweet
Mark Chappell Peter E. Hillman Shawn M. Robinson Ryan Taylor
Charles J. Cole Andrew Holmes Caroline S. Rogers Frank Tiegler
Alison Cree Osamu Kishida Robert Rothman Hugh Tyndale-Biscoe
Chris Crippen Harvey B. Lillywhite Timothy B. Rowe Wayne van Devender
Mark Dimmitt Jessie Maisano Michael J. Ryan John K. VanDyk
Vladimir Dinets Patrick Moldowan Sue Sabrowski David B. Wake
Colleen Farmer Andrew Moore Allen Salzberg Cliff White
Vincent Fernandez Kenneth Nagy Colin Sanders Mark Witton
Luciano Fischer Darren Naish Jay Savage Stephen Zozaya
Linda Ford Gavin Naylor Kurt Schwenk

We leaned especially heavily on many colleagues, and thank them for their generosity:
Robin M. Andrews, Virginia Tech
James Aparicio, Museo Nacional de Historia Natural
de Bolivia
Paul M. Barratt, Natural History Museum, London
Robin M. D. Beck, University of Salford
David R. Begun, University of Toronto
William E. Bemis, Cornell University
Michael J. Benton, University of Bristol
Christopher Brochu, University of Iowa
Edmund D. Brodie, Jr., Utah State University
William S. Brown, Skidmore College
Emily A. Buchholtz, Wellesley College
Ann C. Burke, Wesleyan University
Kenneth Catania, Vanderbilt University
Jennifer A. Clack, University of Cambridge
René C. Clark, Tucson
Chris Crippen, Virginia Living Museum
Michael I. Coates, University of Chicago
A. W. “Fuzz” Crompton, Harvard University
Martha L. Crump, Utah State University
David Cundall, Lehigh University
John D. Damuth, University of California, Santa Barbara
Christine Dahlin, University of Pittsburgh at Johnstown
Dominique Didier, Millersville University
Kenneth Dodd, University of Florida
Phillip C. J. Donoghue, University of Bristol
William N. Eschmeyer, California Academy of Sciences
Robert E. Espinoza, California State University,
Northridge
David E. Fastovsky, University of Rhode Island
Sharon Gilman, Coastal Carolina University
Pamela G. Gill, University of Bristol
Frank M. Greco, New York Aquarium
Gordon Grigg, The University of Queensland
David F. Gruber, City University of New York
Célio F. B. Haddad, Universidade Estadual Paulista “Júlio
de Mesquita Filho”
Lindsay Hazely, The Southland Museum and Art Gallery
Axel Hernandez, Università di Corsica Pasquale Paoli
Thomas R. Holtz, University of Maryland
Carlos Jared, Instituto Butantan
Bruce C. Jayne, University of Cincinnati
Jeffrey Lang, University of North Dakota
Manuel S. Leal, University of Missouri
Agustín Martinelli, Museo Argentino de Ciencias
Naturales “Bernardino Rivadavia”
Phillip Motta, University of South Florida
Sterling J. Nesbit, Virginia Tech
8  CHAPTER 1 Evolution, Diversity, and Classification of Vertebrates
contrast, the age of a fossil can almost
always be determined—provided that
one has a fossil. The difficulty with
fossils is the incompleteness of the
fossil record. There are gaps, some-
times millions of years long, in the
records of many taxa. For example, Turtles Phytosaurs Crocodylians Pterosaurs
we have fossil lampreys from the
Late Devonian (~360 million years
ago, abbreviated Ma1), the Late Car-
boniferous (~300 Ma), and the Early
Cretaceous (~145 Ma), but nothing
between those dates.
Considering the difficulties with
both molecular and morphological
phylogenies, it’s remarkable that they
mostly agree about branching pat-
terns. Disagreements usually center on dates of divergence.
For example, molecular phylogenies indicate that the extant
lineages of amphibians diverged in the Late Carboniferous
(~315–300 Ma), whereas fossil data indicate divergence in
the Late Permian (~260–255 Ma). However, molecular and
morphological methods can agree; molecular evidence in-
dicates that humans separated from their common ancestor
with chimpanzees about 6.6 Ma, and this date fits well
with the earliest fossil in the human lineage, Ardipithecus
(5.8 Ma).
The best information comes from combining molecular
and morphological data. Studies that include extant and
extinct organisms often employ the technique of molecular
scaffolding: the extant taxa are placed in their phylogenetic
position by the relationships established by the molecular
data, and then morphological data are used to integrate the
fossil taxa with the extant taxa. Pough Vertebrate Life 10E Crocodylians are the basal taxon in this tree (the lineage
Sinauer Associates
Morales Studio
Using phylogenetic trees VL10E_01.03.ai 06-1-17rived (farthest from the ancestral form). Together, crocody-
Phylogenetic systematics is based on the assumption that
organisms in a lineage share a common heritage, which
accounts for their similarities. Because of that common
heritage, we can use phylogenetic tress to ask questions
about evolution. By examining the origin and significance
of characters of extant animals, we can make inferences
about the biology of extinct species. For example, the phy-
logenetic relationship of crocodylians, dinosaurs, and birds
is shown in Figure 1.3.
We know that both crocodylians and birds display ex-
tensive parental care of their eggs and young. Some fossil-
ized dinosaur nests contain remains of partly grown baby
dinosaurs, suggesting that at least some dinosaurs cared
for their young. Is that a plausible inference? Obviously,
there is no direct way to determine what sort of parental
care dinosaurs provided to their eggs and young. The inter-
mediate lineages in the phylogenetic tree (pterosaurs and
1
Ma is the abbreviation for mega-annums,“million years.” It is used to
mean both “million years” and, when referring to a specific date range,
“million years ago.”Analogously, ka = kilo-annums (thousand years,
thousand years ago).
Archosauria
Dinosauria
Saurischia
Aves
Ornithischian Saurischian
dinosaurs dinosaurs Birds
Most recent common ancestor
of crocodylians and birds (archosaurs)
Figure 1.3 Using a phylogenetic tree to make inferences about
behavior. The tree shows the relationships of the Archosauria,
the evolutionary lineage that includes today’s crocodylians and
birds. (Phytosaurs were crocodile-like animals that disappeared
at the end of the Triassic; pterosaurs were flying reptiles of the
Jurassic and Cretaceous.) Both extant archosaur groups (i.e., croc-
odylians and birds) display extensive parental care of eggs and
young, but turtles do not. The most parsimonious explanation of
this situation assumes that parental care evolved in the archosaur
lineage after the separation from turtles, but before the separation
of phytosaurs + crocodylians from pterosaurs + dinosaurs.
dinosaurs) are extinct, so we cannot observe their repro-
ductive behavior. However, the phylogenetic tree in Figure
1.3 provides an indirect way to approach the question by
examining the lineage that includes the closest living rel-
atives of dinosaurs, crocodylians, and birds.
closest to the ancestral form), and birds are the most de-
lians and birds form what is called an extant phylogenetic
bracket. Both crocodylians and birds provide parental care.
Looking at extant representatives of more distantly relat-
ed lineages (outgroups), we see that parental care is not
universal among fishes, amphibians, or turtles. The most
parsimonious explanation of the occurrence of parental care
in both crocodylians and birds is that it evolved in that
lineage after the lineage separated from turtles and before
crocodylians separated from dinosaurs + birds. We cannot
prove that parental care did not evolve independently in
crocodylians and in birds, but a single change to parental
care is more likely than two separate changes. Thus, the
most parsimonious hypothesis is that parental care is a
derived character of the evolutionary lineage containing
crocodylians + pterosaurs + dinosaurs + birds. That means
we are probably correct when we interpret fossil evidence
as showing that dinosaurs did indeed exhibit parental care.
Figure 1.3 also shows how phylogenetics has made
talking about restricted groups of animals more compli-
cated than it used to be. Suppose you wanted to refer to just
the two lineages of animals that are popularly known as
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Title: It might have been worse

a motor trip from coast to coast

Author: Beatrice Larned Massey

Release date: September 19, 2023 [eBook #71685]

Language: English

Original publication: San Francisco: Harr Wagner Publishing Co,

1920

Credits: Peter Becker and the Online Distributed Proofreading

Team at https://www.pgdp.net (This file was produced
from images generously made available by The
Internet Archive)

*** START OF THE PROJECT GUTENBERG EBOOK IT MIGHT

HAVE BEEN WORSE ***
 Transcriber’s Note
A much larger, higher-resolution version of the map may
be seen by clicking or right-clicking "(Larger)" beneath it.
Additional notes will be found near the end of this ebook.
 (Larger)
IT MIGHT HAVE BEEN WORSE
KEY
~~~~~~~ ROUTE FOLLOWED BY CAR
- - - - CAR SHIPPED BY BOAT OR TRAIN
—o— TOWNS VISITED, OR EN ROUTE
☉ OVER NIGHT STOPS
it might have been worse
 IT
MIGHT HAVE BEEN
WORSE
A MOTOR TRIP FROM COAST
TO COAST
BY
BEATRICE LARNED MASSEY

SAN FRANCISCO
HARR WAGNER PUBLISHING CO.
MCMXX
Copyright, 1920, By Beatrice Larned Massey
Printed by Taylor & Taylor, San Francisco
TO MY DEAR MR. NIP
 CONTENTS

chapter page
i. the start 1
ii. new york to pittsburgh 6
iii. ohio and detours 20
iv. on to chicago 30
v. through the dairy country 39
vi. clothes, luggage, and the car 43
vii. the twin cities and ten thousand
lakes 54
viii. millions of grasshoppers 62
ix. the bad lands—“nature’s
freakiest mood” 70
x. the dust of montana 77
xi. a wonderland 87
xii. westward ho! 103
xiii. nevada and the desert 117
xiv. the end of the road 130
 FOREWORD

May I state, at the start, that this account of our motor trip from
New York City to San Francisco is intended to be not only a road
map and a motor guide for prospective tourists, but also to interest
the would-be or near motorists who take dream trips to the Pacific? It
sounds like a rather large order, to motor across this vast continent,
but in reality it is simple, and the most interesting trip I have ever
taken in our own country or abroad.
There are so many so-called “highways” to follow, and numerous
routes which, according to the folders, have “good roads and first-
class accommodations all the way” that hundreds of unsuspecting
citizens are touring across every year. I can speak only for
ourselves, and will doubtless call down the criticism of many who
have taken any other route. On the whole, it has been a revelation,
and, to my mind, the only way to get a first-hand knowledge of our
country, its people, the scenery, and last, but not the least, its roads
good, bad, and infinitely worse.
B. L. M.
San Francisco, January, 1920
IT MIGHT HAVE BEEN WORSE
 IT
MIGHT HAVE BEEN
WORSE

THE START

After reading “By Motor to the Golden Gate,” by Emily Post,

published in 1916, I was fired by a desire to make a similar tour. This
desire grew into a firm determination the more I re-read her
charming book. Then the United States went into the war, and self-
respecting citizens were not spending months amusing themselves;
so all thought of the trip was put aside until the spring of this year
(1919). Then the “motor fever” came on again, and refused to yield
to any sedatives of advice or obstacles. After talking and planning for
three years, we actually decided to go in ten minutes—and in ten
days we were off. All the necessary arrangements were quickly
made; leasing our home, storing our household goods, closing up
business matters, getting our equipment and having the car
thoroughly looked over, and all the pleasant but unnecessary duties
occupied the last few days. Why will people write so many letters
and say so many good-bys, when a more or less efficient mail and
telegraph service circles our continent? But it is the custom, and all
your friends expect it—like sending Easter and Christmas cards by
the hundreds. We are victims of a well-prescribed custom.
It is always of interest to me to know the make of car that a
friend (or stranger) is driving; so let me say, without any desire to
advertise the Packard, that we had a new twin-six touring car, of
which I shall speak later on. I believe in giving just tribute to any car
that will come out whole and in excellent condition, without any
engine troubles or having to be repaired, after a trip of 4154 miles
over plains and mountains, through ditches, ruts, sand, and mud,
fording streams and two days of desert-going. And let me add that
my husband and I drove every mile of the way. It is needless to say
that the car was not overstrained or abused, and was given every
care on the trip. In each large city the Packard service station
greased and oiled the car, turned down the grease-cups, examined
the brakes and steering-gear, and started us off in “apple-pie” order,
with a feeling on our parts of security and satisfaction.
The subject of car equipment, tires, clothes, and luggage will
take a chapter by itself. But let me say that we profited in all these
regards by the experience and valuable suggestions of Mrs. Post in
her book.
When we first spoke to our friends of making this trip, it created
as little surprise or comment as if we had said, “We are going to tour
the Berkshires.” The motor mind has so grown and changed in a few
years. Nearly everyone had some valuable suggestion to make, but
one only which we accepted and profited by. Every last friend and
relative that we had offered to go in some capacity—private
secretaries, chauffeurs, valets, maids, and traveling companions. But
our conscience smote us when we looked at that tonneau, the size
of a small boat, empty, save for our luggage, which, let me add with
infinite pride and satisfaction, was not on the running-boards, nor
strapped to the back. From the exterior appearance of the car we
might have been shopping on Fifth Avenue.
We extended an invitation to two friends to accompany us, which
was accepted by return mail, with the remark, “Go!—of course, we
will go! Never give such an invitation to this family unless you are in
earnest.” And so our genial friends joined us, and we picked them up
at the Seymour Hotel in New York City, at three o’clock, Saturday,
July 19th, and started for the Forty-second-Street ferry in a pouring
rain, as jolly and happy a quartette as the weather would permit. Our
guests were a retired physician, whom we shall speak of as the
Doctor, and his charming, somewhat younger wife, who, although
possessing the perfectly good name of Helen, was promptly dubbed
“Toodles” for no reason in the world. These dear people were of the
much-traveled type, who took everything in perfect good-nature and
were never at all fussy nor disturbed by late hours, delays, bad
weather, nor any of the usual fate of motorists, and they both added
to the pleasure of the trip as far as they accompanied us.
It had rained steadily for three days before we started and it
poured torrents for three days after; but that was to be expected, and
the New Jersey and Pennsylvania roads were none the worse, and
the freedom from dust was a boon. We chose for the slogan of our
trip, “It might have been worse.” The Doctor had an endless fund of
good stories, of two classes, “table and stable stories,” and I regret
to say that this apt slogan was taken from one of his choicest stable
stories, and quite unfit for publication. However, it did fit our party in
its optimism and cheery atmosphere.
With a last look at the wonderful sky-line of the city, and the hum
and whirl of the great throbbing metropolis, lessening in the swirl of
the Hudson River, we really were started; with our faces turned to
the setting sun, and the vast, wonderful West before us.
 II

NEW YORK TO PITTSBURGH

One of the all-absorbing pleasures in contemplating a long trip is

to map out your route. You hear how all your friends have gone, or
their friends, then you load up with maps and folders, especially
those published by all the auto firms and tire companies, you pore
over the Blue Book of the current year, and generally end by going
the way you want to go, through the cities where you have friends or
special interests. This is exactly what we did. As the trip was to be
taken in mid-summer, we concluded to take a northern route from
Chicago, via Milwaukee, St. Paul, Fargo, Billings, Yellowstone Park,
Salt Lake City, Ogden, Reno, Sacramento, to San Francisco (see
map), and, strange to relate, we followed out the tour as we had
planned it. With the exception of a few hot days in the larger cities
and on the plains, and, of course, in the desert, we justified our
decision.
As I have stated, we drove 4154 miles, through sixteen states
and the Yellowstone Park, in thirty-three running days, and the trip
took just seven weeks to the day, including seventeen days spent in
various cities, where we rested and enjoyed the sights. As time was
of no special object, and we were not attempting to break any
records, we felt free to start and stop when we felt inclined to do so;
on only two mornings did we start before nine-thirty, and seldom
drove later than seven in the evening. In so doing, we made a
pleasure of the trip and not a duty, and avoided any unusual fatigue.
The first evening we reached Easton, Pennsylvania. We were
glad to get into the comfortable Huntington Hotel out of the wet, and
enjoyed a good dinner and a night’s rest. We followed the Lincoln
Highway to Pittsburgh, and have only praise to offer for the condition
of the road and the beauty of the small towns through which we
went. Of all the states that we crossed, Pennsylvania stands out par
excellence in good roads, clean, attractive towns, beautiful farming
country and fruit belts, and well-built, up-to-date farm buildings. In
other states we found many such farms, but in Pennsylvania it was
exceptional to find a poor, tumble-down farmhouse or barn. The
whole state had an air of thrift and prosperity, and every little home
was surrounded by fine trees, flowers, and a well-kept vegetable
garden.
The worst bugbear of the motorist are the detours. Just why the
road commissioners choose the height of the motoring season to
tear up the main highways and work the roads has always been a
mystery to me, and I have never heard any logical solution of it. We
were often told that no work to speak of had been done on the state
roads through the country during the war, and in many places the
heavy army trucks had cut up the good roads until the ruts left turtle-
backed ridges in the center, not at all pleasant to bob along on. But,
in view of what we encountered later in our trip, I look back on the
Pennsylvania roads as one of the high spots and pleasures, never to
be undervalued.
From Easton we drove in the rain to Harrisburg. The scenery
was beautiful. The Blue Ridge and the Alleghany Mountains loomed
up in the haze like great cathedrals; but as long as the road was
wide and comparatively smooth we enjoyed the ups and downs. Our
engine told us that we were gradually ascending; the mist would be
wafted off by a mountain breeze, and then a gorgeous panorama
stretched before us as far as the eye could see.
We found Harrisburg a busy, thriving city, with well-paved
streets, attractive homes, and many fine buildings. The leading hotel,
the Penn Harris, was turning away guests; so we were made very
comfortable at the Senate. Here the café was miserable, but we
went to the restaurant of the Penn Harris and had an excellent
dinner at moderate prices. We have found that at the largest, best
hotels the food was better cooked and much cheaper than at the
smaller ones. Usually we had excellent club breakfasts from forty