Vertebrate Palaeontology and Evolution

711 Pages·43.593 MB·English

by  Carroll R.L.

VERTEBRATE
PALEONTOLOGY
AND
EVOLUTION
ROBERT L. CARROLL
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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
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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