Vertebrate Palaeontology

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

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.