Michael J. Benton, OBE, FRS, is Professor of Vertebrate Palaeontology at the University of Bristol. He is particularly interested in early reptiles, Triassic dinosaurs and macroevolution, and has published over 60 books and 700 scientific articles. He is part of one of the most active palaeontology research groups in the world and has supervised more than 80 PhD students.

List of Boxes xvii

Preface xxi

About the Companion Website xxv

1 Vertebrates Originate 1

Introduction 1

1.1 Sea Squirts and the Lancelet 2

1.2 Ambulacraria: Echinoderms and Hemichordates 4

1.3 Deuterostome Relationships 6

1.4 Chordate Origins 9

1.5 Vertebrate Origins: Worm or Bag? 15

1.6 Further Reading 20

Questions for Future Research 20

References 21

2 How to Study Fossil Vertebrates 25

Introduction 25

2.1 Digging Up Bones 25

2.2 Publication and Professionalism 32

2.3 Geology and Fossil Vertebrates 37

2.4 Biology and Fossil Vertebrates 46

2.5 Discovering Phylogeny 53

2.6 Macroevolution 56

2.7 Further Reading 59

References 59

3 Early Palaeozoic Fishes 63

Introduction 63

3.1 Cambrian Vertebrates 63

3.2 Vertebrate Hard Tissues 65

3.3 The Jawless Fishes 67

3.4 Origin of Jaws 80

3.5 Placoderms: Armour-Plated Monsters 82

3.6 Pan-Chondrichthyes: Sharks, Rays, Acanthodians 90

3.7 Early Fish Biogeography and Environments 92

3.8 Osteichthyes: The Bony Fishes 94

3.9 Early Fish Evolution and Mass Extinction 106

3.10 Further Reading 107

Questions for Future Research 108

References 108

4 The First Tetrapods 115

Introduction 115

4.1 Problems of Life on Land 115

4.2 Devonian Tetrapods 120

4.3 The Carboniferous World 128

4.4 Diversity of Carboniferous Tetrapods 129

4.5 Temnospondyls and Reptiliomorphs After the Carboniferous 139

4.6 Evolution of the Modern Amphibians 144

4.7 Further Reading 150

Questions for Future Research 150

References 150

5 Evolution of Early Amniotes 157

Introduction 157

5.1 Hylonomus and Paleothyris - Biology of the First Amniotes 157

5.2 Amniote Adaptations for Terrestrial Life 160

5.3 The Permian World 167

5.4 The Parareptiles 169

5.5 The Eureptiles 174

5.6 Early Synapsid Evolution 176

5.7 The Permian-Triassic Mass Extinction 188

5.8 Further Reading 192

Questions for Future Research 192

References 193

6 The Triassic Revolution 197

Introduction 197

6.1 The Triassic World and the Recovery of Life 197

6.2 Triassic Marine Reptiles 202

6.3 Evolution of the Archosauromorphs 211

6.4 Origin of the Dinosaurs 220

6.5 Amniote Evolution in the Triassic 224

6.6 Further Reading 235

Questions for Future Research 235

References 236

7 Evolution of Fishes After the Devonian 243

Introduction 243

7.1 The Early Chimaeras and Sharks 243

7.2 Post-Palaeozoic Chondrichthyan Radiation 251

7.3 The Early Bony Fishes 255

7.4 Radiation of the Teleosts 267

7.5 Post-Devonian Evolution of Fishes 276

7.6 Further Reading 279

Questions for Future Research 279

References 280

8 Dinosaurs 287

Introduction 287

8.1 Biology of Plateosaurus 287

8.2 The Jurassic and Cretaceous World 289

8.3 Saurischians and Theropod Diversity 290

8.4 The Sauropodomorph Dinosaurs 303

8.5 The Diversity of Ornithischian Dinosaurs 309

8.6 Were the Dinosaurs Warm-Blooded or Not? 326

8.7 Further Reading 333

Questions for Future Research 334

References 334

9 The Mesozoic Reptiles 343

Introduction 343

9.1 Testudinata: The Turtles 343

9.2 Crocodylomorpha 353

9.3 Pterosauria 359

9.4 The Great Sea Dragons 367

9.5 Lepidosauria: Lizards and Snakes 374

9.6 The End-Cretaceous Mass Extinction 383

9.7 Further Reading 390

Questions for Future Research 390

References 390

10 The Birds 401

Introduction 401

10.1 The Origin of Birds 401

10.2 The Origin of Bird Flight 412

10.3 Cretaceous Birds, With and Without Teeth 418

10.4 The Radiation of Modern Birds: Explosion or Long Fuse? 429

10.5 Flightless Birds: Palaeognathae 432

10.6 Neognathae 435

10.7 The Three-Phase Diversification of Birds 447

10.8 Further Reading 448

Questions for Future Research 448

References 449

11 Mammals: Origins and Southern Hemisphere Evolution 459

Introduction 459

11.1 Cynodonts and the Acquisition of Mammaliaform Characters 459

11.2 The First Mammaliaforms 472

11.3 The Mesozoic Mammaliaforms 475

11.4 Marsupials Down Under 492

11.5 South American Mammals - A World Apart 497

11.6 Afrotheria and the Break-up of Gondwana 506

11.7 Further Reading 512

Questions for Future Research 513

References 513

12 Mammals of the Northern Hemisphere 525

Introduction 525

12.1 Evolution of Modern Mammals 525

12.2 Boreoeutherian Beginnings: The Palaeocene in the Northern Hemisphere 529

12.3 Early Diverging Laurasiatherians: Eulipotyphla 537

12.4 Scrotifera: Bats and Relatives 540

12.5 Cetartiodactyla: Cattle, Pigs and Whales 540

12.6 Zooamata: Horses, Carnivores, and Pangolins 550

12.7 Glires: Rodents, Rabbits, and Relatives 563

12.8 Archonta: Primates, Tree Shrews, and Flying Lemurs 568

12.9 Ice Age Extinction of Large Mammals 569

12.10 Further Reading 574

Questions for Future Research 574

References 574

13 Human Evolution 585

Introduction 585

13.1 What Are the Primates? 585

13.2 The Fossil Record of Early Primates 586

13.3 Anthropoidea: Monkeys and Apes 592

13.4 Hominoidea: The Apes 597

13.5 Evolution of Human Characteristics 603

13.6 The Early Stages of Human Evolution 605

13.7 The Past Two Million Years of Human Evolution 611

13.8 Further Reading 620

Questions for Future Research 620

References 620

Appendix: Classification of the Vertebrates 629

Glossary 641

Index 647