VERTEBRATE PALAEONTOLOGY Dedicated to Mary, Philippa and Donald for their forebearance while I wrote this book. VERTEBRATE PALAEONTOLOGY Fourth edition MICHAEL J. BENTON School of Earth Sciences University of Bristol Bristol, UK Chapter opening illustrations drawn by John Sibbick This edition first published 2015 © 2015 Michael J. Benton Third edition published 2005 © 2005 Blackwell Publishing Ltd. Second edition published 1997 by Chapman & Hall. First edition published 1990 by Unwin Hyman. Registered Office John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial Offices 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 111 River Street, Hoboken, NJ 07030-5774, USA For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley-blackwell. The right of the author to be identified as the author of this work has been asserted in accordance with the UK Copyright, Designs and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher. Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. Limit of Liability/Disclaimer of Warranty: While the publisher and author(s) have used their best efforts in preparing this book, they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose. It is sold on the understanding that the publisher is not engaged in rendering professional services and neither the publisher nor the author shall be liable for damages arising herefrom. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloging-in-Publication Data Benton, M. J. (Michael J.) Vertebrate palaeontology / Michael J. Benton. – Fourth edition. pages cm Includes bibliographical references and index. ISBN 978-1-118-40755-4 (cloth) – ISBN 978-1-118-40684-7 (pbk.) 1. Vertebrates, Fossil. I. Title. QE841.B44 2015 566–dc23 2014000064 A catalogue record for this book is available from the British Library. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books. Cover image: Lobster lunch of Luoping; over a muddy seabed during the Middle Triassic (Ansian) at the future site of Luoping County, Yunnan. In the foreground, an early rock lobster (Yunnanopalinura schrami) attracts the unwanted attention of the coelacanth Luopingcoelacanthus eurylacrimalis and the armoured reptile Sinosaurosphargis yunguiensis. Behind them cruise other aquatic reptiles: the ichthyosaurs Mixosaurus cf. panxianensis and the little pachypleurosaur Dianopachysaurus dingi. Swarming around is a diverse community of actinopterygians: ubiquitous Sangiorgioichthys sui (numerous fish with black-tipped fins), sleek Sinosaurichthys longimedialis (thin with a long snout), tiny Marcopoloichthys ani (very small with red-spotted fins), scaleless Gymnoichthys inopinatus (blue with a yellow back) and the deep-bodied Kyphosichthys grandei (brown and yellow) and Luoxiongichthys hyperdorsalis (black and white). Painting by Brian Choo © 2013. Set in 9.5/12pt Minion by SPi Publisher Services, Pondicherry, India 1 2015 Contents LIST OF BOXES viii 3.1 Cambrian vertebrates 46 PREFACE x 3.2 Vertebrate hard tissues 49 ABOUT THE COMPANION WEBSITE xii 3.3 The jawless fishes 51 3.4 Origin of jaws and gnathostome relationships 59 3.5 Placoderms: armour-plated monsters 60 1 VERTEBRATES ORIGINATE 1 3.6 Chondrichthyes: the first sharks 65 3.7 Acanthodians: the ‘spiny skins’ 65 Key questions in this chapter 2 3.8 Devonian environments 67 Introduction 2 3.9 Osteichthyes: the bony fishes 70 1.1 Sea squirts and the lancelet 2 3.10 Early fish evolution and mass extinction 79 1.2 Ambulacraria: echinoderms and hemichordates 4 3.11 Further reading 79 1.3 Deuterostome relationships 6 Questions for future research 80 1.4 Chordate origins 8 3.12 References 80 1.5 Vertebrates and the head 14 1.6 Further reading 14 Questions for future research 15 4 EARLY TETRAPODS AND AMPHIBIANS 84 1.7 References 15 Key questions in this chapter 85 Introduction 85 2 HOW TO STUDY FOSSIL VERTEBRATES 18 4.1 Problems of life on land 85 4.2 Devonian tetrapods 88 Key questions in this chapter 19 4.3 The Carboniferous world 96 Introduction 19 4.4 Diversity of Carboniferous tetrapods 98 2.1 Digging up bones 19 4.5 Temnospondyls and reptiliomorphs after 2.2 Publication and professionalism 24 the Carboniferous 106 2.3 Geology and fossil vertebrates 29 4.6 Evolution of the modern amphibians 109 2.4 Biology and fossil vertebrates 33 4.7 Further reading 114 2.5 Discovering phylogeny 36 Questions for future research 114 2.6 The quality of the fossil record 39 4.8 References 114 2.7 Macroevolution 41 2.8 Further reading 43 2.9 References 43 5 EvOLUTION OF EARLY AMNIOTES 118 3 EARLY PALAEOZOIC FISHES 45 Key questions in this chapter 119 Introduction 119 Key questions in this chapter 46 5.1 Hylonomus and Paleothyris – biology Introduction 46 of the first amniotes 119 vi Contents 5.2 Amniote evolution 121 8.11 The Cretaceous-Paleogene mass extinction 259 5.3 The Permian world 125 8.12 Further reading 263 5.4 The parareptiles 125 Questions for future research 264 5.5 The eureptiles 128 8.13 References 264 5.6 Basal synapsid evolution 132 5.7 The Permo-Triassic mass extinction 142 5.8 Further reading 143 9 THE BIRDS 273 Questions for future research 143 5.9 References 143 Key questions in this chapter 274 Introduction 274 9.1 The origin of birds 274 6 BOUNCEBACK: TETRAPODS OF THE TRIASSIC 147 9.2 The origin of bird flight 282 9.3 Cretaceous birds, with and without teeth 287 Key questions in this chapter 148 9.4 The radiation of modern birds: explosion Introduction 148 or long fuse? 296 6.1 The Triassic world and its effect on 9.5 Flightless birds: Palaeognathae 299 the recovery of life 148 9.6 Neognathae 300 6.2 Triassic marine reptiles 150 9.7 The three-phase diversification of birds 311 6.3 Evolution of the archosauromorphs 154 9.8 Further reading 312 6.4 Origin of the dinosaurs 161 Questions for future research 313 6.5 Reptile evolution in the Triassic 164 9.9 References 313 6.6 Further reading 168 Questions for future research 168 6.7 References 168 10 MAMMALS 318 Key questions in this chapter 319 7 EvOLUTION OF FISHES AFTER Introduction 319 the deVONIAN 172 10.1 Cynodonts and the acquisition of mammalian characters 319 Key questions in this chapter 173 10.2 The first mammals 328 Introduction 173 10.3 The Mesozoic mammals 332 7.1 The early sharks and chimaeras 173 10.4 Evolution of modern mammals 343 7.2 Post-Palaeozoic chondrichthyan radiation 178 10.5 Marsupials down under 346 7.3 The early bony fishes 181 10.6 South American mammals – a world apart 349 7.4 Radiation of the teleosts 191 10.7 Afrotheria and the break-up of Gondwana 355 7.5 Post-Devonian evolution of fishes 198 10.8 Boreoeutherian beginnings: the Palaeocene 7.6 Further reading 199 in the northern hemisphere 361 Questions for future research 199 10.9 Basal laurasiatherians: Lipotyphla 366 7.7 References 199 10.10 Cetartiodactyla: cattle, pigs and whales 366 10.11 Pegasoferae: bats, horses, carnivores and pangolins 375 8 THE AGE OF DINOSAURS 204 10.12 Glires: rodents, rabbits and relatives 383 10.13 Archonta: primates, tree shrews and Key questions in this chapter 205 flying lemurs 388 Introduction 205 10.14 Ice age extinction of large mammals 389 8.1 Biology of Plateosaurus 205 10.15 Further reading 390 8.2 The Jurassic and Cretaceous world 206 Questions for future research 390 8.3 The diversity of saurischian dinosaurs 207 10.16 References 391 8.4 The diversity of ornithischian dinosaurs 221 8.5 Were the dinosaurs warm-blooded or not? 232 8.6 Pterosauria 236 11 HUMAN EVOLUTION 400 8.7 Testudinata: the turtles 241 8.8 Crocodylomorpha 247 Key questions in this chapter 401 8.9 Lepidosauria: lizards and snakes 250 Introduction 401 8.10 The great sea dragons 256 11.1 What are the primates? 401 ____________________________________________________________________________________________ Contents vii 11.2 The fossil record of early primates 402 APPENDIX: CLASSIFICATION OF 11.3 Anthropoidea: monkeys and apes 407 THE VERTEBRATES 433 11.4 Hominoidea: the apes 411 11.5 Evolution of human characteristics 414 GLOSSARY 448 11.6 The early stages of human evolution 416 INDEX 453 11.7 The past two million years of human evolution 421 11.8 Further reading 428 Questions for future research 428 The colour plate section can be found between pp. 244–245 11.9 References 429 Preface To many, palaeontology in general, and vertebrate palaeontol- of the international appeal of vertebrate palaeontology and ogy in particular, might be seen as devoted to discovering new the demand from students and instructors for up-to-date fossils. After all, we read lavish press reports of each new species information.