This page intentionally left blank From the authors of The Evolution and Extinction of the Dinosaurs comes a new textbook designed to excite undergraduates about science by using dinosaurs to illustrate and discuss geology, natural history, and evolution. Emphasizing the logic of science over facts and details, the fundamental concepts of dinosaurs – origins – diversity – behavior – extinction – are conveyed in concise, lively text with exceptional and unique illustrations. Hypothesis testing and scientifi c concepts drive the strong narrative. Students are introduced to novel and revolutionary ideas on the natural history of dinosaurs; ideas that will likely change their perception of the biota and their place in it. Fastovsky and Weishampel root the text in the common language of modern evolutionary biology – phylogenetic systematics – requiring students to assess data critically, like all practicing scientists. Gideon Mantell (1790–1852), the “father” of modern dinosaur paleontology. Dinosaurs A Concise Natural History David E. Fastovsky University of Rhode Island and David B. Weishampel The Johns Hopkins University With illustrations by John Sibbick CAMBRIDGE UNIVERSITY PRESS Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo Cambridge University Press The Edinburgh Building, Cambridge CB2 8RU, UK Published in the United States of America by Cambridge University Press, New York www.cambridge.org Information on this title: www.cambridge.org/9780521889964 © Cambridge University Press 2009 This publication is in copyright. Subject to statutory exception and to the provision of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published in print format 2009 ISBN-13 978-0-511-47941-0 eBook (EBL) ISBN-13 978-0-521-88996-4 hardback ISBN-13 978-0-521-71902-5 paperback Cambridge University Press has no responsibility for the persistence or accuracy of urls for external or third-party internet websites referred to in this publication, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate. Contents Why a natural history of dinosaurs? ix Summary 45 To the student ix Appendix 3.1: What is “evolution”? 46 To the instructor x Selected readings 48 Dedication xiii Topic questions 48 Part I: Reaching back in time 1 1 To catch a dinosaur 3 4 Who are the dinosaurs? 51 Chapter objectives 51 Chapter objectives 3 Finding the history of life 52 Tales of dinosaurs 4 In the beginning 52 Fossils 4 Tetrapoda 55 Collecting 9 Box 4.1: Fish and chips 58 Summary 16 Box 4.2: What, if anything, is a “reptile”? 62 Selected readings 17 Diapsida 63 Topic questions 17 Dinosaurs 65 Origins 67 2 Dinosaur days 18 Box 4.3: Stance: it’s both who you are and Chapter objectives 18 what you do 68 When did dinosaurs live and how do we Ornithischia and Saurischia 70 know? 20 Summary 71 Continents and climates 24 Selected readings 72 Climates during the time of the dinosaurs 27 Topic questions 73 Summary 29 Appendix 2.1: Chemistry quick ’n dirty 29 Part II: Ornithischia: armored, horned, and d uck-billed Selected readings 30 dinosaurs 75 Topic questions 31 What makes an ornithischian an ornithischian? 76 3 Who’s related to whom – and how do we Chew on this! 77 know? 32 Ornithischia: the big picture 79 Chapter objectives 32 Selected readings 83 Who are you? 34 Evolution 34 5 Thyreophorans: the armor-bearers 85 Phylogenetic systematics – the reconstruction of Chapter objectives 85 phylogeny 36 Thyreophora 86 Box 3.1: Wristwatches: when is a watch a Eurypoda: Stegosauria – hot plates 87 watch? 42 Box 5.1: The poetry of dinosaurs 92 Science and testing hypotheses 45 vi Contents Box 5.2: Dino brains 93 Prosauropoda 163 Eurypoda: Ankylosauria – mass and gas 95 Sauropoda 165 The evolution of Thyreophora 102 Box 8.1: Every breath you take 172 Summary 105 The evolution of Sauropodomorpha 175 Selected readings 106 B ox 8.2: The recapitation of “Brontosaurus” 179 Topic questions 107 Summary 183 Selected readings 184 6 M arginocephalia: bumps, bosses, and beaks 108 Topic questions 184 Chapter objectives 109 9 Theropoda I: nature red in tooth and claw 187 Marginocephalia 110 Chapter objectives 187 Marginocephalia: Pachycephalosauria – In Domes We Trust 110 Theropoda 188 The evolution of Pachycephalosauria 117 Theropod lives and lifestyles 189 Marginocephalia: Ceratopsia – horns and all the Box 9.1: Triceratops spoils or spoiled frills 118 Triceratops? 202 The evolution of Ceratopsia 128 The evolution of Theropoda 205 Summary 130 Summary 209 Selected readings 132 Selected readings 209 Topic questions 133 Topic questions 211 7 Ornithopoda: the tuskers, antelopes, and 10 Theropoda II: the origin of birds 213 “mighty ducks” of the Mesozoic 135 Chapter objectives 213 Birds 214 Chapter objectives 135 Archaeopteryxand the ancestry of living birds 217 Ornithopoda 136 Box 10.1: Plus ça change . . . 225 Box 7.1: Hypotheses that didn’t fl oat 139 Feathers without fl ight 226 The evolution of Ornithopoda 149 Summary 231 Summary 152 Selected readings 232 Selected readings 152 Topic questions 233 Topic questions 153 Part III: Saurischia: meat, might, and 11 Theropoda III: early birds 235 magnitude 155 Chapter objectives 235 Mesozoic birds 236 Saurischia: the big picture 156 Evolution of Aves 240 Selected readings 159 Box 11.1: Molecular evolution and the origin of Aves 241 8 Sauropodomorpha: the big, the bizarre, and the Summary 243 majestic 161 Selected readings 244 Chapter objectives 161 Topic questions 244 Sauropodomorpha 162 Contents vii Part IV: Endothermy, endemism, and Box 14.3: Dinosaur wars in the nineteenth extinction 247 century: boxer versus puncher 298 Box 14.4: Louis Dollo and the beasts of 12 Dinosaur thermoregulation: some like it hot 248 Bernissart 301 Chapter objectives 249 The second part of the twentieth century to today 303 The way they were 250 Box 14.5: Rollin’ on the river 304 Temperature talk 250 Box 14.6: “Mr Bones” 306 Dinosaur endothermy: the evidence 250 Box 14.7: Tendaguru! 307 Box 12.1: The “skinny” on metabolism 251 Box 14.8: Baron Franz von Nopcsa: Box 12.2: Warm-bloodedness: to have and to nationalism, Transylvanian dinosaurs, and have hot 252 espionage 309 Box 12.3: In the tracks of dinosaurs 254 Summary 315 Box 12.4: Dinosaur smarts 256 Box 14.9: Young Turks and old turkeys 316 Box 12.5: Weighing in 263 Selected readings 318 Summary 267 Topic questions 319 Selected readings 268 Topic questions 269 15 The Cretaceous–Tertiary extinction: the frill is gone 321 13 The fl owering of the Mesozoic 271 Chapter objectives 321 Chapter objectives 271 How important were the deaths of a few dino- Dinosaurs in the Mesozoic Era 274 saurs? 322 B ox 13.1: The shape of tetrapod diversity 278 Geological record of the latest Cretaceous 322 Box 13.2: Counting dinosaurs 280 Biological record of the latest Cretaceous 327 Plants and dinosaurian herbivores 282 Box 15.1: Extinction 329 Box 13.3: Dinosaurs invent fl owering Box 15.2: Dinosaurs: all wrong for mass plants 287 extinctions 335 Summary 287 Extinction hypotheses 336 Selected readings 288 Box 15.3: The real reason the dinosaurs Topic questions 289 became extinct 340 Summary 342 14 A history of paleontology through ideas 291 Selected readings 343 Chapter objectives 291 Topic questions 344 The idea of ideas 292 In the beginning 292 Glossary 345 Box 14.1: Indiana Jones and the Central Figure credits 361 Asiatic Expedition of the American Museum Index of subjects 363 of Natural History 293 Index of genera 374 The nineteenth century through the mid- twentieth century 295 Box 14.2: Sir Richard Owen: brilliance and darkness 296