Table Of Content

Introduction to Paleobiology and the Fossil Record Companion website This book includes a companion website at: www.blackwellpublishing.com/paleobiology The website includes: (cid:129) An ongoing database of additional Practicals prepared by the authors (cid:129) Figures from the text for downloading (cid:129) Useful links for each chapter (cid:129) Updates from the authors Introduction to Paleobiology and the Fossil Record Michael J. Benton University of Bristol, UK David A. T. Harper University of Copenhagen, Denmark A John Wiley & Sons, Ltd., Publication This edition fi rst published 2009, © 2009 by Michael J. Benton and David A.T. Harper Blackwell Publishing was acquired by John Wiley & Sons in February 2007. Blackwell’s publishing program has been merged with Wiley’s global Scientifi c, Technical and Medical business to form Wiley-Blackwell. Registered offi ce: John Wiley & Sons Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial offi ces: 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 offi ces, 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 identifi ed as the author of this work has been asserted in accordance with the 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. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books. 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. This publication is designed to provide accurate and authoritative information in regard to the subject matter covered. It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloguing-in-Publication Data Benton, M. J. (Michael J.) Introduction to paleobiology and the fossil record / Michael J Benton, David A.T. Harper. p. cm. Includes bibliographical references and index. ISBN 978-1-4051-8646-9 (hardback : alk. paper) – ISBN 978-1-4051-4157-4 (pbk. : alk. paper) 1. Evolutionary paleobiology. 2. Paleobiology. 3. Paleontology. I. Harper, D. A. T. II. Title. QE721.2.E85B46 2008 560–dc22 2008015534 A catalogue record for this book is available from the British Library. Set in 11 on 12 pt Sabon by SNP Best-set Typesetter Ltd, Hong Kong Printed in Singapore by Markono Print Media Pte Ltd 1 2009 Contents Full contents vii Preface xi 1 Paleontology as a science 1 2 Fossils in time and space 22 3 Taphonomy and the quality of the fossil record 57 4 Paleoecology and paleoclimates 79 5 Macroevolution and the tree of life 116 6 Fossil form and function 137 7 Mass extinctions and biodiversity loss 162 8 The origin of life 183 9 Protists 204 10 Origin of the metazoans 234 11 The basal metazoans: sponges and corals 260 12 Spiralians 1: lophophorates 297 13 Spiralians 2: mollusks 326 14 Ecdysozoa: arthropods 361 15 Deuterostomes: echinoderms and hemichordates 389 16 Fishes and basal tetrapods 427 17 Dinosaurs and mammals 453 18 Fossil plants 479 19 Trace fossils 509 20 Diversifi cation of life 533 Glossary 554 Appendix 1: Stratigraphic chart 573 Appendix 2: Paleogeographic maps 575 Index 576 A companion resources website for this book is available at http://www.blackwellpublishing.com/paleobiology Full contents Preface xi 1 Paleontology as a science 1 Paleontology in the modern world 2 Paleontology as a science 3 Steps to understanding 9 Fossils and evolution 12 Paleontology today 13 Review questions 20 Further reading 20 References 21 2 Fossils in time and space 22 Frameworks 23 On the ground: lithostratigraphy 25 Use of fossils: discovery of biostratigraphy 25 Paleobiogeography 41 Fossils in fold belts 48 Review questions 55 Further reading 55 References 55 3 Taphonomy and the quality of the fossil record 57 Fossil preservation 58 Quality of the fossil record 70 Review questions 77 Further reading 77 References 78 4 Paleoecology and paleoclimates 79 Paleoecology 80 Paleoclimates 103 Review questions 113 Further reading 113 References 114 5 Macroevolution and the tree of life 116 Evolution by natural selection 118 Evolution and the fossil record 120 viii FULL CONTENTS The tree of life 128 Review questions 135 Further reading 136 References 136 6 Fossil form and function 137 Growth and form 138 Evolution and development 144 Interpreting the function of fossils 150 Review questions 159 Further reading 160 References 160 7 Mass extinctions and biodiversity loss 162 Mass extinctions 163 The “big fi ve” mass extinction events 169 Extinction then and now 178 Review questions 181 Further reading 181 References 181 8 The origin of life 183 The origin of life 184 Evidence for the origin of life 188 Life diversifi es: eukaryotes 195 Review questions 202 Further reading 202 References 202 9 Protists 204 Protista: introduction 206 Eukaryotes arrive center stage 207 Protozoa 208 Chromista 226 Review questions 232 Further reading 233 References 233 10 Origin of the metazoans 234 Origins and classifi cation 235 Four key faunas 241 Soft-bodied invertebrates 256 Review questions 257 Further reading 257 References 257 11 The basal metazoans: sponges and corals 260 Porifera 261 Cnidaria 271 Review questions 296 Further reading 296 References 296 FULL CONTENTS ix 12 Spiralians 1: lophophorates 297 Brachiopoda 298 Bryozoa 313 Review questions 324 Further reading 324 References 324 13 Spiralians 2: mollusks 326 Mollusks: introduction 327 Early mollusks 327 Class Bivalvia 332 Class Gastropoda 338 Class Cephalopoda 344 Class Scaphopoda 354 Class Rostroconcha 354 Evolutionary trends within the Mollusca 355 Review questions 360 Further reading 360 References 360 14 Ecdysozoa: arthropods 361 Arthropods: introduction 362 Early arthropod faunas 362 Subphylum Trilobitomorpha 363 Subphylum Chelicerata 375 Subphylum Myriapoda 379 Subphylum Hexapoda 381 Subphylum Crustacea 381 Review questions 387 Further reading 387 References 387 15 Deuterostomes: echinoderms and hemichordates 389 Echinoderms 390 Hemichordates 409 Review questions 425 Further reading 425 References 425 16 Fishes and basal tetrapods 427 Origin of the vertebrates 428 Jaws and fi sh evolution 435 Tetrapods 442 Reign of the reptiles 443 Review questions 451 Further reading 451 References 451 17 Dinosaurs and mammals 453 Dinosaurs and their kin 454 Bird evolution 460