VERTEBRATE PALEONTOLOGY AND EVOLUTION ROBERT L. CARROLL ALSO AVAILABLE FROM W.H. FREEMAN AND COMPANY EARTH AND LIFE THROUGH TIME Steven M. Stanley "A real tour de force." -Charles W. Pitrat, University of Massachusetts This acclaimed text stresses the interrelationshíps between geology and biology, emphasizing the effects that such events as plate movements, climate change, sea-level fluctuations, and mountain growth and destruction have on one another; how these events determine the direction in which life evolves; and how living things, in turn, have changed the face of the Earth. 1986, cloth, 690 pages, 897 iJlustrations ISBN 0-7167-1677-1 / EXTINCTION Steven M. Stanley This lavishly illustrated, full-color book examines the cataclysmic outbreaks of mass extinction that have eliminated tens of thousands of thriving species throughout the evolutionary record. Stanley focuses on what the rich storehouse of rock and fossil clues can teli us about these dramatíc global disasters-how they originated and progressed, and what they imply for the future. 1987, cloth, 242 pages, 183 illustrations, 134 ín color ISBN 0-7167-5014-7 FOSSILS AND THE HISTORY OF LIFE George Gaylord Simpson "A beautiful, well written book." - James 0. Farlow, Indiana University-Purdue University at Fort Wayne ln the late George Gaylord Simpson's )ast published work, the internationally known paleontologist examines the rich, worldwide fossil record upon which many of the theories and facts of evolution are based, and he offers his persona! view of paleontology and its relevance for understanding the history of life. 1984, cloth, 239 pages, 165 illustrations ISBN 0-7167-1564-3 FIVE K.JNGDOMS An Illustrated Guide to the Phyla of Life on Earth Second Edition Lynn Margulis and Karlene V. Schwartz '. "lt is remarkable that no one had previously thought of producing such a comprehensive, obvious, and valuable document." - from the Foreword by Stephen Jay Gould Five Kingdoms is a profusely illustrated guide to the 92 phyla of the five kingdoms that make up all life on Earth. lt provides an accurate, up-to-date reference for science students, instructors, and anyone else who is curious about the extraordinary variety of living things that inhabit this plaoet. 1987, cloth, 416 pages, 421 illustrations ISBN 0-7167-1885-5 ISBN 0-7167-1822-7 Vertebrate Paleontology and Evolution is the first detailed, authoritative review of the fos sil record of vertebrates to be published in 20 years. Distinguished paleontologist Robert L. Carroll provides complete, current data on all fossil groups, which he examines in the con text of evolutionary processes and functional anatomy. The skeletal structure and way of life of the major groups are discussed, as are their interrelationships and distribution in time and space. This integrated approach provides a remarkably coherent analysis of the history of vertebrates through 600 million years of evolution. This extraordinary new work features more than 1700 illustrations, including 500 newly drafted line drawings of the represen tative members of all major vertebrate groups. Forty phylogenies and cladograms depict relationships among major groups. Significant discoveries and the formulation of new theories over the past two decades The Author have vastly increased our knowledge of fossil vertebrates, especially of earlier forms from Robert L. Carroll is professor of biology at the Paleozoic and Mesozoic. Newly dis McGill University, Montreal, and is director of covered forms of jawless fish, sharks, and that institution's Redpath Museum, where he amphibians are described here in full detail, has been curator of vertebrate paleontology as are the major transitions between amphibi since 1965. He graduated from Michigan State ans and reptiles, reptiles and mammals, and University (B.S.) and Harvard University (M.S., dinosaurs and birds. Ph.D.), wherehestudied under A. S. Romer. ln Carroll pays particular attention to sub 1978 he received the Schuchert Award of the jects in which conflicting evidence has led to Paleontological Society in recognition of his alternative hypotheses, such as the origin of excellence in paleontology. He was president of vertebrates, the emergence of each of the the Society ofVertebrae Paleontology in 1983. major groups, and the cataclysmic extinction Dr. Carroll's major research interests are the at the end of the Mesozoic. He also demon anatomy and relationships of Paleozoic and strates the importance of functional anatomy early Mesozoic amphibians and reptiles, and in providing a deeper understanding of the the origin of the modem amphibian and reptile nature and relationships of extinct organ groups. He has published numerous technical isms, and he comments at length on the papers on fossil vertebrates from the western resurgent concern over the methodology of United States, eastern Canada, Madagascar, classification. southern Africa, Great Britain, central Europe, Vertebrate Paleontology and Evolution is a and China. ln addition to his many duties at completely modern, up-to-date, and indis McGill and the Redpath Museum, Dr. Carroll pensable volume for all professional verte sits on the executive board of the First World brate paleontologists and students of geo Congress of Herpetology and is associate editor logy, biology, and evolutionary theory. of the Canadian ]ournal of Earth Sciences. "1 am greatly impressed by the data pre Cover photograph courtesy of Staatliches sented ... Useful not only for students but for Museum für Naturkunde in Stuttgart. colleagues in the field as a resource." Hans-Peter Schultze, University of Kansas "A real tour de force: clearly written; com 11 prehensive and detailed without slipping into needless jargon; well-researched, up-to-date, and scholarly." W. H. FREEMAN AND COMPANY 41 Madison Avenue, New York, NY 10010 Léo Laporte, University of California, 20 Beaumont Street, Oxford OXl 2NQ, Santa Cruz England VERTEBRATE PALEONTOLOGY AND EVOLUTION ROBERT L. CARROLL Redpath Museum McGill University W.H. FREEMAN AND COMPANY NEW YORK Líbrary of Congress Cataloging-in-Publícation Data Carroll, Robert L Vertebrate paleontology and evolution. Includes index. 1. Vertebrates, Fossil. 2. Vertebrates-Evolution. !. Title. QE84 LC254 1987 5 66 86-31808 ISBN 0-716-71822-7 Copyright © 1988 by W. H. Freeman and Company No part of this book may be reproduced by any mechanícal, photographic, or electronic process, or in the form of a phonographic recording, nor may it be stored ín a retrieval system, transmitted, or otherwise copíed for public or privatc use without written permission from the publisher. Printed ín the United States of America. 34567890 HL 6543210898 Dedicated to the memory of my father, John H. Carro11 who introduced me to the joys of paleonrology e s 0 N T E N T Preface Xlll CHAPTER I Fossils and Relarionships 1 The Nature of the Fossil Record 2 Evolution 4 Establishing Relationships 5 Polarity 8 Cladograms 10 Paraphyly and Holophyly 11 Summary 13 CHAPTER II The Origin of Vertebrates 16 The Body Plan of Primitív e Vertebrates 17 Invertebrate q Chordates 18 The Origin of Vcrtebrate Characters 21 viií CONTENTS 1 The Origin of Skeletal CHAPTER VII Advanced Bony Tissues 22 Fish: The Teleosts 112 The Environment of T eleost Classification 113 Early Vertebrates 23 Ancestral T eleosts 113 Summary 24 Living Teleost Groups 115 Euteleosts 118 Neoteleosts 120 CHAPTER I1I The Diversity of Jawless Fish 26 Spiny Teleosts- The Acanthomorpha 122 Ostracoderms 26 Primitive Living Jawless Acanthomorphs- Vertebrates 39 The Ctenothrissoids 124 lnterrelationships Paracanthopterygians 125 of Primitive Acanthopterygians 126 Vertebrates 41 Summary 133 Summary 42 CHAPTER VIII Sarcopterygian Fish 136 CHAPTER IV Primitív e The Anatomy of jawed Fish 44 Lobe-finned Fish 136 The Origin of Jaws 44 Rhi pidistians 138 Placoderms 46 Struniiformes 146 Arthrodira 49 Coelacanthiformes Antiarchi 52 (Actinistia) 147 Phyllolepidida 53 Lungfish 148 Petalichthyida 54 Alternative Ptyctodontida 54 Relationships 153 Acanthothoraci 55 Summary 154 Rhenanida 56 Stensioellidae and CHAPTER IX The Conquest of Paraplesiobatidae 58 Land and the Interrelationships Radiation of of Placoderms 58 Amphibians 156 Summary 58 Paleozoic Amphibians 158 The Origin of CHAPTER V Sharks and Other Amphibians 166 Cartilaginous Fish 62 The Radiation Advanced of Paleozoic Jawed Fish 62 Amphibians 167 Paleozoic Modern Amphibians 180 Elasmobranchs 63 Summary 188 Neoselachii 74 Holocephali 78 CHAPTER X Primitive Amniotes Summary 82 and Turtles 192 Relationships of CHAPTER Vl Acanthodians and Amniotes 193 Primitive Bony Fish 84 Primitive Amniotes 193 Acanthodians 85 Temporal Openings Osteichthyes 89 and the Actinopterygians 90 Classification of Primitív e Amniotes 199 Actinopterygians: The Diversity among Palaeonisciformes 90 Primitive Anapsids 201 Diversity of Testudines 207 Chondrostean Fish 98 Summary 214 Neopterygian Characteristics 102 CHAPTER XI Primitive Diapsids Prímitive Neopterygians 107 and Lepidosaurs 217 Summary 110 Early Diapsids 217 CONTENTS ix 1 Characteristics of CHAPTER XVII The Origin of Advanced Diapsids 220 Mammmals 361 Eosuchians 221 Pelycosaurs 362 Gephyrosaurus and Therapsíds 369 Champsosaurs 224 The Origin of Lepidosaurs 224 the Mammalian Summary 238 Skeleton 392 Summary 397 CHAPTER XII Mesozoic Marine Reptiles 240 CHAPTER XVIII Primitive Mesozoic Sauropterygians 241 Mammals and Placodonts 248 Monotremes 401 lchthyosaurs 251 The Anatomy of Míddle and Upper Morganucodon 402 Triassic The Bio\ogy of Ichthyosaurs 254 Primitive Mammals 410 Jurassic and The Diversity of Cretaceous Primitive Mammals 413 lchthyosaurs 256 Interrelationships of Summary 258 Mesozoic Mammals 419 Monotremes 420 CHAPTER xm Primitive Summary 422 Archosauromorphs and Crocodiles 261 CHAPTER XIX Primitive Therian Primitive Mammals and Archosauromorphs 264 Marsupials 425 Thecodontia 268 The Evolution of Crocodylia 277 Therian Molar Summary 283 Teeth 425 Marsupials 430 CHAPTER XIV The Anatomy and Summary 439 Relationships of Dinosaurs 286 CHAPTER XX The Radiation of The Posture and Placental Mammals 443 Ancestry of Placenta! Dinosaurs 288 Reproduction 443 Saurischians 289 Lower Cretaceous Theropods 289 Eutherians 445 Sauropodomorphs 297 Upper Cretaceous Ornithischians 302 Eutherians 445 Segnosauria 314 The Beginning of the Summary 315 Cenozic Mammalian CHAPTER XV The Biology and Radiation 449 Extinction of Geography 453 Dinosaurs 319 Eutherian Mammals Biology 320 lncertae Sedis 454 Birth and Growth 325 T aeniodonta 456 The Extinction at the Tillodonts and End of the Pantodonts 457 Mesozoic 325 Dinocerata (Uintatheres) 459 CHAPTER XVI Flight 331 Insectivora 460 Pterosaurs 331 Bats 463 Mesozoic Birds 338 Primates 464 Cenozoic Birds 344 Scandentia Summary 357 (Tree Shrews) 477 x: CONTENTS Dermoptera 477 Edentates 554 Carnivores 478 The Great Interchange 563 Rodents, Lagomorphs, Summary 563 and Elephant Shrews 485 CHAPTER XXII Evolution 569 Summary 495 Evolution at the Species Levei 569 CHAPTER XXI Ungulates, Edentates, Macroevol ution 576 and Whales 502 Constraínts and Condylarths 502 Radiatíons 581 Artiodactyla 507 Extinction 587 Mesonychia 520 Summary 590 Cetaceans 522 Perissodactyla 527 APPENDIX Classification of African Ungulates 536 Vertebrates 594 South American Ungulates 547 Index 649
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