David E. Fastovsky University of Rhode Island David B. Weishampel Johns Hopkins University With original illustrations by Brian Regal, Tarbosaurus Studio A'gJ" CAMBRIDGE UNIVERSITY PRESS
Preface xv CHAPTER 1 CHAPTER 2 INTRODUCTION FOSSILS 5 Taphonomy 5 COLLECTION 11 Prospecting 12 Collecting 13 Back at the Ranch 16 PARTI SETTING THE STAGE THE MESOZOIC ERA: BACK TO THE PAST WHEN DID DINOSAURS LIVE AND HOW DO WE KNOW? 20 Chronostratigraphy 21 Age in Years 21 Lithostratigraphy 25 Biostratigraphy 26 Eras and Periods and Epochs, Oh My! 27 Growth of a Prehistoric Time Scale 29 PLATE TECTONICS: A DRIVING FORCE IN EVOLUTION? 29 Structure of the Earth 30 Motions of the Plates 30 Plates through Time 33 CLIMATES THROUGH THE MESOZOIC ERA 38 Potential Effects of Plate Motion on Climate 38 Summary of Climates through the Mesozoic 43 Box 2.1 Stable Isotopes, Ancient Temperatures, and Dead Oceans 1 3 19 44 CHAPTER 3 DISCOVERING ORDER IN THE NATURAL WORLD HIERARCHY 49 Characters 50 Cladograms 51 49 Vll
vm CHAPTER 4 ORGANIC EVOLUTION 54 Evolution 54 Cladograms and the Reconstruction of Phylogeny 57 Parsimony 60 Box 3.1 The Evolution of Wrist Watches Science and Testing Hypotheses 63 INTERRELATIONSHIPS OF THE VERTEBRATES THE EARLY BIOTA AND ITS HISTORY 66 Body Plans 69 Box 4.1 The Biological Classification The Middle Cambrian Explosion and the Earliest Vertebrates 71 CHORDATES 73 Vertebrata 75 Gnathostomata 75 TETRAPODA 76 The Tetrapod Skeleton 77 Amniota 82 Box 4.2 Posture: It's Both Who You Are and What Tou Do Synapsida 84 REPTILIA 86 Anapsid 86 Diapsida 86 Archosauria 87 Dinosaurs 89 Box 4.3 Fish and Chips 62 66 70 82 90 CHAPTER 5 THE ORIGIN OF DINOSAURIA HISTORY OF DINOSAURIA 94 Dinosaurs - The Mesozoic Unnaturals 95 Dinosauria - It's a Natural! 97 ORIGINS 99 THE RISE OF DINOSAURS: SUPERIORITY OR LUCK? 101 94 PART II ORNITHISCHIA ORNITHISCHIA: ARMORED, HORNED, AND DUCKBILLED DINOSAURS 107 109 CHAPTER 6 STEGOSAURIA: HOTPLATES HISTORY OF STEGOSAUR DISCOVERIES 114 113
IX 19th Century 114 20th Century 115 STEGOSAURIA DEFINED AND DIAGNOSED 119 STEGOSAUR DIVERSITY AND PHYLOGENY 119 STEGOSAUR PALEOBIOLOGY AND PALEOECOLOGY: GAITS, PATES, AND PLATES 122 Stance and Gait 123 Dealing with Mealing 123 Box 6.1 19th-century Dinosaur Wars: Boxer vs. Puncher 124 Brains 126 Brains between the Thighs 127 Box 6.2 The Poetry of Dinosaurs 128 Behavior 129 \ Plates and Spines 129 CHAPTER 7 ANKYLOSAURIA: MASS AND GAS 134 HISTORY OF ANKYLOSAUR DISCOVERIES 135 19th Century 135 20th Century 136 ANKYLOSAURIA DEFINED AND DIAGNOSED 139 ANKYLOSAUR DIVERSITY AND PHYLOGENY: ACTORS IN THE ANKYLOSAUR ARMORY 139 Box 7.1 Indiana Jones and the Central Asiatic Expeditions of the American Museum of Natural History 140 ANKYLOSAUR PALEOBIOLOGY AND PALEOECOLOGY: FRISKY, RUNNING, LEAPING ANKYLOSAURS - NOT! 146 Food and Mouths to Feed 146 Defensive Moves 148 CHAPTER 8 PACHYCEPHALOSAURIA: HEAD-TO-HEAD, WITH MALICE AFORETHOUGHT 151 HISTORY OF PACHYCEPHALOSAUR DISCOVERIES 151 19th Century 151 20th Century 152 PACHYCEPHALOSAURIA DEFINED AND DIAGNOSED 156 PACHYCEPHALOSAUR DIVERSITY AND PHYLOGENY 157 PACHYCEPHALOSAUR PALEOBIOLOGY AND PALEOECOLOGY: FOOD AND SEX 160 Butting 162 CHAPTER 9 CERATOPSIA: HORNS, FRILLS, AND SLICE-AND-DICE 169 HISTORY OF CERATOPSIAN DISCOVERIES 172 19th Century 172
CHAPTER 10 20th Century 172 CERATOPSIA DEFINED AND DIAGNOSED 175 CERATOPSIAN DIVERSITY AND PHYLOGENY 176 Protoceratopsidae... Or Not? 177 Ceratopsidae 179 CERATOPSIAN PALEOBIOLOGY AND PALEOECOLOGY: FEEDING, LOCOMOTION, HORNS, AND ALL THE FRILLS 182 Feeding 182 Locomotion 185 Socializing the Dinosaur 185 ORNITHOPODA: THE TUSKERS, ANTELOPES, AND THE MIGHTY DUCKS OF THE MESOZOIC HISTORY OF ORNITHOPODA DISCOVERIES 191 19th Century 191 20th Century 194 ORNITHOPODA DEFINED AND DIAGNOSED 198 ORNITHOPOD DIVERSITY AND PHYLOGENY 199 Heterodontosauridae 199 Euornithopoda 199 Hypsilophodontidae 202 Iguanodontia 203 Hadrosauridae 204 ORNITHOPOD PALEOBIOLOGY AND PALEOECOLOGY 209 Geography 209 Habits 209 Feeding and Food 212 Box 10.1 Swimming in the Treetops and Other Dormant Hypotheses Head Structures and Behavior 216 Family Values in the Mesozoic: Bringing Up Baby 220 190 214 PART III SAURISCHIA SAURISCHIA: PREDATORS AND GIANTS OF THE MESOZOIC 225 226 CHAPTER 11 SAUROPODOMORPHA: THE BIG, THE BIZARRE, AND THE MAJESTIC HISTORY OF THE STUDY OF SAUROPODOMORPHA 230 19th Century 230 20th Century 232 229
XI Box 11.1 Tendagurul SAUROPODOMORPHA DEFINED AND DIAGNOSED 238 SAUROPODOMORPH DIVERSITY AND PHYLOGENY 238 Sauropoda 240 Camarasaurs and Titanosaurs 242 SAUROPODOMORPH PALEOBIOLOGY AND PALEOECOLOGY: LIFESTYLES OF THE HUGE AND ANCIENT 246 Distribution and Preservation 246 Habits 247 Box 11.2 Brontosaurus: Beheading a Wrong-Headed Dinosaur Feeding 254 Locomotion 256 Social Behavior 258 Sauropodomorph Growth and Development 258 234 250 CHAPTER 12 THEROPODA I: NATURE RED IN TOOTH AND CLAW HISTORY OF THEROPOD DISCOVERIES 263 19th Century 263 20th Century 263 THEROPODA DEFINED AND DIAGNOSED 271 THEROPOD DIVERSITY AND PHYLOGENY 272 Basal Theropods? 272 Ceratosauria 272 Tetanurae 275 Above Maniraptora 276 Dromaeosaurids and Their Kin 278 THEROPOD PALEOECOLOGY AND PALEOBIOLOGY 279 Theropods as Living Organisms 281 Box 12.1 Triceratops as Spoils or Spoiled Triceratopsi 261 285 CHAPTER 13 THEROPODA II: THE ORIGIN OF BIRDS WHAT IS A BIRD? 294 ARCHAEOPTERYX LITHOGRAPHICA AND THE ANCESTRY OF BIRDS 298 Anatomy of Archaeopteryx 300 Birds as Dinosaurs 304 Pneumatic Bones and Feathers: Adaptation vs. Inheritance 309 Flight 311 THE EARLY EVOLUTION OF AVES 314 Getting to B.e a Modern Bird 314 Early Cretaceous 314 Late Cretaceous Birds 317 293
Xll CHAPTER 14 PARTIV ENDOTHERMY, ENVIRONMENTS, AND EXTINCTION DINOSAUR ENDOTHERMY: SOME LIKE IT HOT THE GENESIS OF AN IDEA 325 The Therapsid-Dinosaur Transition 325 ENDOTHERMY AND ECTOTHERMY 329 Temperature Regulation among Vertebrates 329 Box 14.1 To Have and to Have Hot Endothermic and Ectothermic Metabolism 331 DINOSAUR ENDOTHERMY: THE EVIDENCE 333 Anatomy 333 Box 14.2 In the Tracks of the Dinosaurs Histology 340 Box 14.3 Dinosaur Smarts Box 14.4 Weighing in a Dinosaur Ecology 345 Zoogeography 349 Phylogeny 352 Geochemistry 353 The Nose Knows 354 TOWARD A CONSENSUS? 355 y 323 325 329 336 340 344 CHAPTER 15 DINOSAURS IN SPACE AND TIME GEOGRAPHIC DISTRIBUTION 360 Box 15.1 Counting Dinosaurs TRIASSIC-JURASSIC 363 Late Triassic (231-208 Ma) 363 Early Jurassic (208-178 Ma) 366 Middle Jurassic (178-157Ma) 369 Box 15.2 The Shape oftetrapod Diversity Late Jurassic (157-145 Ma) 372 CRETACEOUS 374 Early Cretaceous (145-97 Ma) 374 Late Cretaceous (97-65 Ma) 377 Box 15.3 Dinosaurs Invent Flowering Plants THE TERTIARY: AFTER THE BALL IS OVER 385 359 362 370 381
CHAPTER 16 CHAPTER 17 RECONSTRUCTING EXTINCTIONS: THE ART OF SCIENCE EXTINCTIONS 388 RESOLVING THE PAST 390 Incompleteness of the Fossil Record: Resolution 390 Range Extensions 394 Choice of Taxonomic Level and the Problem of Incongruent Taxa 395 THE CRETACEOUS-TERTIARY EXTINCTION: THE FRILL IS GONE GEOLOGICAL RECORD OF THE LATEST CRETACEOUS 400 Tectonic Activity 400 Oceans 401 Climate 401 Asteroid Impact 401 Volcanism 407 Smoking Gun 407 BIOLOGICAL RECORD OF THE LATEST CRETACEOUS 411 Oceans 411 Terrestrial Record 414 THE EXTINCTION HYPOTHESES 420 Extinction Hypotheses 420 Box 17.1 The Real Reason the Dinosaurs Became Extinct xin 387 399 424 COLOR PLATES after 366 GLOSSARY 431 SUBJECT INDEX 443 TAXONOMIC INDEX OF GENERA 453 AUTHOR INDEX 458