Vertebrate Palaeontology

Michael J. Benton 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 50 books and 300 scientific articles. He leads one of the most successful palaeontology research groups at the University of Bristol, and has supervised over 60 PhD students.

List of boxes viii Preface x

About the companion website xii

1 Vertebrates Originate 1

Key questions in this chapter 2

Introduction 2

1.1 Sea squirts and the lancelet 2

1.2 Ambulacraria: echinoderms and hemichordates 4

1.3 Deuterostome relationships 6

1.4 Chordate origins 8

1.5 Vertebrates and the head 14

1.6 Further reading 14

Questions for future research 15

1.7 References 15

2 How to Study Fossil Vertebrates 18

Key questions in this chapter 19

Introduction 19

2.1 Digging up bones 19

2.2 Publication and professionalism 24

2.3 Geology and fossil vertebrates 29

2.4 Biology and fossil vertebrates 33

2.5 Discovering phylogeny 36

2.6 The quality of the fossil record 39

2.7 Macroevolution 41

2.8 Further reading 43

2.9 References 43

3 Early palaeozoic fishes 45

Key questions in this chapter 46

Introduction 46

3.1 cambrian vertebrates 46

3.2 vertebrate hard tissues 49

3.3 the jawless fishes 51

3.4 origin of jaws and gnathostome relationships 59

3.5 placoderms: armour-plated monsters 60

3.6 chondrichthyes: the first sharks 65

3.7 acanthodians: the ‘spiny skins’ 65

3.8 devonian environments 67

3.9 osteichthyes: the bony fishes 70

3.10 early fish evolution and mass extinction 79

3.11 further reading 79

Questions for future research 80

3.12 References 80

4 Early tetrapods and amphibians 84

Key questions in this chapter 85

Introduction 85

4.1 Problems of life on land 85

4.2 Devonian tetrapods 88

4.3 The Carboniferous world 96

4.4 Diversity of Carboniferous tetrapods 98

4.5 Temnospondyls and reptiliomorphs after the Carboniferous 106

4.6 E volution of the modern amphibians 109

4.7 Further reading 114

Questions for future research 114

4.8 References 114

5 Evolution of early amniotes 118

Key questions in this chapter 119

Introduction 119

5.1 Hylonomus and Paleothyris – biology of the first amniotes 119

5.2 Amniote evolution 121

5.3 The Permian world 125

5.4 The parareptiles 125

5.5 The eureptiles 128

5.6 Basal synapsid evolution 132

5.7 The Permo-Triassic mass extinction 142

5.8 Further reading 143

Questions for future research 143

5.9 References 143

6 Bounceback: tetrapods of the Triassic 147

Key questions in this chapter 148

Introduction 148

6.1 The Triassic world and its effect on the recovery of life 148

6.2 Triassic marine reptiles 150

6.3 E volution of the archosauromorphs 154

6.4 O rigin of the dinosaurs 161

6.5 Reptile evolution in the Triassic 164

6.6 Further reading 168

Questions for future research 168

6.7 References 168

7 Evolution of fishes after the devonian 172

Key questions in this chapter 173

Introduction 173

7.1 The early sharks and chimaeras 173

7.2 Post-Palaeozoic chondrichthyan radiation 178

7.3 The early bony fishes 181

7.4 Radiation of the teleosts 191

7.5 Post-Devonian evolution of fishes 198

7.6 Further reading 199

Questions for future research 199

7.7 References 199

8 The age of dinosaurs 204

Key questions in this chapter 205

Introduction 205

8.1 Biology of Plateosaurus 205

8.2 The Jurassic and Cretaceous world 206

8.3 The diversity of saurischian dinosaurs 207

8.4 The diversity of ornithischian dinosaurs 221

8.5 Were the dinosaurs warm-blooded or not? 232

8.6 Pterosauria 236

8.7 Testudinata: the turtles 241

8.8 Crocodylomorpha 247

8.9 Lepidosauria: lizards and snakes 250

8.10 The great sea dragons 256

8.11 The Cretaceous-Paleogene mass extinction 259

8.12 Further reading 263

Questions for future research 264

8.13 References 264

9 The birds 273

Key questions in this chapter 274

Introduction 274

9.1 The origin of birds 274

9.2 The origin of bird flight 282

9.3 Cretaceous birds, with and without teeth 287

9.4 The radiation of modern birds: explosion or long fuse? 296

9.5 Flightless birds: palaeognathae 299

9.6 Neognathae 300

9.7 The three-phase diversification of birds 311

9.8 Further reading 312

Questions for future research 313

9.9 References 313

10 Mammals 318

Key questions in this chapter 319

Introduction 319

10.1 Cynodonts and the acquisition of mammalian characters 319

10.2 The first mammals 328

10.3 The Mesozoic mammals 332

10.4 Evolution of modern mammals 343

10.5 Marsupials down under 346

10.6 South American mammals – a world apart 349

10.7 Afrotheria and the break-up of Gondwana 355

10.8 Boreoeutherian beginnings: the Palaeocene in the northern hemisphere 361

10.9 Basal laurasiatherians: Lipotyphla 366

10.10 Cetartiodactyla: cattle, pigs and whales 366

10.11 Pegasoferae: bats, horses, carnivores and pangolins 375

10.12 Glires: rodents, rabbits and relatives 383

10.13 Archonta: primates, tree shrews and flying lemurs 388

10.14 Ice age extinction of large mammals 389

10.15 Further reading 390

Questions for future research 390

10.16 References 391

11 Human evolution 400

Key questions in this chapter 401

Introduction 401

11.1 What are the primates? 401

11.2 The fossil record of early primates 402

11.3 Anthropoidea: monkeys and apes 407

11.4 Hominoidea: the apes 411

11.5 E volution of human characteristics 414

11.6 The early stages of human evolution 416

11.7 The past two million years of human evolution 421

11.8 Further reading 428

Questions for future research 428

11.9 References 429

Appendix: Classification of the vertebrates 433

Glossary 448

Index 453

The colour plate section can be found between pp. 244–245