VERTEBRATE PALEONTOLOGY AND EVOLUTION Mee CCC a og Swe, 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 interrelationships 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 illustrations ISBN 0-7167-1677-1 EXTINCTION Steven M. Stanley This lavishly illustrated, fall-color book examines the cataclysmic outbreaks of mass extinction that have climinated tens of thousands of thriving species throughout the evolutionary record. Stanley focuses on what the rich storchouse of rock and fossil clues can tell us about these dramatic global disasters—how they originated and progressed, and what they imply for the future. 1987, cloth, 242 pages, 183 illustrations, 134 in color ISBN 0-7167-5014-7 FOSSILS AND THE HISTORY OF LIFE George Gaylord Simpson “A beautiful, well written book.” —James O. Farlow, Indiana University—Purdue University at Fort Wayne In the late George Gaylord Simpson’s last 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 personal view of paleontology and its relevance for understanding the history of life 1984, cloth, 239 pages, 165 illustrations ISBN 0-7167-1564-3 FIVE KINGDOMS An Illustrated Guide to the Phyla of Life on Earth Second Edition Lynn Margulis and Karlene V. Schwartz “It is remarkable that ao one had previously thought of producing such a comprehensive, obvious, and valuable document.” —from the Foreword by Stephen Jay Gould Five Kingdomsis a profusely illustrated guide to the 92 phyla ofthe five kingdoms that make up all life on Earth. It 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 chis planet. 1987, cloth, 416 pages, 421 illustrations ISBN 0-7167-1885-5 ISBN 0-7167-1822-7Vertebrate Paleontology and Evolution isthe 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 have vastly increased our knowledge of fossil vertebrates, especially of earlier forms from the Paleozoic and Mesozoic. Newly dis- covered forms of jawless fish, sharks, and amphibians are described here in full detail, as are the major transitions between amphibi- ans and reptiles, reptiles and mammals, and dinosaurs and birds. Carroll pays particular attention to sub- jects in which conflicting evidence has led to alternative hypotheses, such as the origin of vertebrates, the emergence of each of the major groups, and the cataclysmic extinction at the end of the Mesozoic. He also demon- strates the importance of functional artatomy in providing 2 deeper understanding of the nature and relationships of extinct organ- isms, and he comments at length on the resurgent concern over the methodology of classification. Vertebrate Paleontology and Evolution is a completely modern, up-to-date, and indis- pensable volume for all professional verte- brate paleontologists and students of geo- logy, biology, and evolutionary theory. “Lam greatly impressed by the data pre- sented... Useful tot only for students but for colleagues in the field as a resource.” Hans-Peter Schultze, University of Kansas “A real tour de force: clearly written; com- prehensive and detailed without slipping into needless jargon; well-researched, up-to-date, and scholarly.” Léo Laporte, University of California, Santa Cruz ‘The Author Robert L. Carroll is professor of biology at McGill University, Montreal, and is director of that institution's Redpath Museum, where he has been curator of vertebrate paleontology since 1965. He graduated from Michigan State University (B.S.) and Harvard University (M.S., Ph.D.), where he studied under A. S. Romer. In 1978 he received the Schuchert Award of the Paleontological Society in recognition of his excellence in paleontology. He was president of the Society of Vertebrae Paleontology in 1983. Dr. Carroll's major research interests are the anatomy and relationships of Paleozoic and carly Mesozoic amphibians and reptiles, and theorigin of the modern amphibian and reptile ‘groups. He has published numerous technical papers on fossil vertebrates from the western United States, eastern Canada, Madagascar, southern Africa, Great Britain, central Europe, and China, In addition to his many duties at McGill and the Redpath Museuta, Dr. Carroll sits on the executive board of the First World Congress of Herpetology and is associate editor of the Canadian Journal of Earth Sciences. Cover photograph courtesy of Staatliches Museum fisr Naturkunde in Stuttgart, W.H, FREEMAN AND COMPANY 41 Madison Avenue, New York, NY 10010 20 Beaumont Street, Oxford OX1 2NQ, EnglandVERTEBRATE PALEONTOLOGY AND EVOLUTION ROBERT L. CARROLL Redpath Museum McGill University W.H FREEMAN AND COMPANY NEW YORKLibrary of Congress Cataloging-in-Publication Data Carroll, Robert L. Vertebrate paleontology and evolution. Includes index. 1, Vertebrates, Fossil. 2. Vertebrates—Evolution, I, Tile. QES41.C254 1987 56686-31808 ISBN 0-716-71822-7 Copyright © 1988 by W. H. Freeman and Company. No part of this book may be reproduced by any mechanical, photogeaphic, tr electronic process, or in the form of a phonographic recording, nor may it be stored in a retrieval system, transmitted, or otherwise copied for public or private use without written permission from the publisher. Printed in the United States of America. 34567890 HL 6543210898Dedicated to the memory of my father, John H, Carroll who introduced me to the joys of paleontoiogyCc O N T Preface CHAPTER I Fossils and Relationships The Nature of the Fossil Record Evolution Establishing, Relationships Polarity Cladograms Paraphyly and Holophyly Summary CHAPTER I The Origin of Vertebrates The Body Plan of Primitive Vertebrates Invertebrate Chordates The Origin of Vertebrate Characters 18 2CHAPTER Il CHAPTER IV CHAPTER V CHAPTER VI The Origin of Skeletal Tissues ‘The Environment of Early Vertebrates Summary ‘The Diversity of Jawless Fish Ostracoderms Living Jawless Vertebrates Interrelationships of Primitive Vertebrates Summary Primitive Jawed Fish The Origin of Jaws Placoderms Arthrodira Antiarchi Phyllolepidida Petalichthyida Pryctodontida Acanthothoraci Rhenanida Stensioellidae and Paraplesiobatidae Intertelationships of Placoderms Summary Sharks and Other Cartilaginous Fish ‘Advanced Jawed Fish Paleozoic Elasmobranchs Neosdachii Holocephali Summary Acanthodians and Primitive Bony Fish Acanthodians Osteichthyes Actinopterygians Primitive ‘Actinopterygians: The Palaconiscitormes Diversity of ‘Chondrostean Fish Neoprerygian Characteristics Primitive Neopcerygians Summary 2B 24 26 26 39 aL 42 44 44 49 52 53 54 54 35 56 58 58 58 62 62 63 74 78 82 84 85 89 90 90 98 102 17 116 CONTENTS, CHAPTER VIL CHAPTER VII CHAPTER IX CHAPTER X CHAPTER XI ‘Advanced Bony Fish: The Teleosts Teleost Classification Ancestral Teleosts Living Teleost Groups Euteleosts Neoteleosts Spiny Teleosts— ‘The Acamhomorpha Primitive Acanthomorphs— ‘The Ctenothrissoids Paracanthopterygians Acanthopterygians Summary Sarcopterygian Fish The Anatomy of Lobe-finned Fish Rhipidistians Seeuniiformes Coelacanthiformes (Actinistia) Lungfish Alternative Relationships Summary The Conquest of Land and the Radiation of Amphibians Paleozoic Amphibians ‘The Origin of Amphibians The Radiation of Paleozoic Amphibians Modern Amphibians Summary Primitive Amniotes and Turtles Relationships of Amniotes Primitive Amniotes ‘Temporal Openings and the Classification of Amniotes Diversity among Primitive Anapsids ‘Testudines Summary Primitive Diapsids and Lepidosaurs Early Diapsids 124 125 126 133, 136 136 138 146 147 148 153 154 156 158 166 167 180 188 192 193 193 199 201 207 214 217 217CHAPTER Xl CHAPTER XIII CHAPTER XIV CHAPTER XV CHAPTER XVI CONTENTS Characteristics of ‘Advanced Diapsids Eosuchians Gephyrosaurus and Champsosaurs Lepidosaurs Summary ‘Mesozoic Marine Reptiles Sauropterygians Placodonts Ichthyosaurs. Middle and Upper Triassic Ichthyosaurs Jurassic and Cretaceous Ichthyosaurs Summary Primitive Archosauromorphs and Crocodiles Primitive Archosauromorphs ‘Thecodontia Crocodylia Summary ‘The Anatomy and Relationships of Dinosaurs The Posture and Ancestry of Dinosaurs Saurischians Theropods Sauropodomorphs Ornithischians Segnosauria Summary The Biology and Extinction of Dinosaurs Biology Birth and Growth The Extinction at the End of the Mesozoic Flight Pterosaurs: Mesozoic Birds Cenozoic Birds Summary 220 221 224 224 238 240 241 248, 251 254 256 258 261 264 268 277 283 286 288 289 289 297 302 314 315s 319 320 325 325 331 331 338, 344 357 CHAPTER XVII CHAPTER XVII CHAPTER XIX CHAPTER XX The Origin of Mammmals Pelycosaurs ‘Therapsids ‘The Origin of the Mammalian Skeleton Summary Primitive Mesozoic Mammals and Monotremes The Anatomy of Morganucodon ‘The Biology of Primitive Mammals The Diversity of Primitive Mammals Interrelationships of Mesozoic Mammals ‘Monotremes Summary Primitive Therian Mammals and Marsupials ‘The Evolution of Therian Molar Teeth Marsupials Summary The Radiation of Placental Mammals Placental Reproduction Lower Cretaceous Eutherians Upper Cretaceous Eutherians ‘The Beginning of the Mammalian Radiation Geography Eutherian Mammals Incertae Sedis Taeniodonta Tillodonts and Pantodonts Dinocerata (Vintatheres) Insectivora Bats Primates Scandentia (Tree Shrews) 361 362 369) 392 397 401 402 410 413 419 420 422 425 425 430 439 443 443 445 445 449 453 454 456 457 459 460 463 464 47Dermoptera Carnivores Rodents, Lagomorphs, and Elephant Shrews Summary CHAPTER XXt_— Ungulates, Edentates, and Whales Condylarths Artiodactyla Mesonychia Cetaceans Perissodactyla African Ungulates, South American Ungulates 477 478 485 495, 502 502 507 520 $22 527 536 547 | CONTENTS CHAPTER XXI APPENDIX Edentates ‘The Great Interchange Summary Evolution Evolution at the Species Level Macroevolution Constraints and Radiations Extinetion Summary Classification of Vertebrates Index 554 563 563 569 569, 576 581 587 590 594 649‘PRET aT TT AT Last night in the museum's hall The fossils gathered for a ball. There were no drums or saxophones, Bur just the clatter of their bones, A rolling, rattling carefree circus Of mammoth polkas and mazurkas. Prerodactyls and brontosauruses Sang ghostly prehistoric choruses. Amid the mastodonic wassail I caught the eye of one small fossil. Cheer up, sad world, he said, and winked— It’s kind of fun to be extinct. Ogden Nash “Carnival of the Animals”* + Reprinted by permission of Curtis Brown, Ltd. Copyright 1950 by Ogden Nash. Renewed 1978 by Frances Nash.Vertebrate Paleontology and Evolution is intended as a comprehensive text for university courses in vertebrate biology and as a reference for biologists and geologists ‘who want to become familiar with the pattern of verte brate evolution. I hope that it also provides the necessary background for research into more specific details of the anatomy, relationships, and distribution of both fossil and living vertebrates. During the past 20 years, our knowledge of fossil vertebrates has increased immensely. Entirely new groups of jawless fish, sharks, amphibians, and dinosaurs have been discovered, and the major transitions between am- phibians and reptiles, reptiles and mammals, and dino- saurs and birds have been thoroughly studied. Evidence from both paleontology and molecular biology provides much new information on the initial radiation of both birds and placental mammals. This book integrates knowledge of the anatomy of extinct vertebrates to serve as the basis for understanding the origin and relationships of the modern orders. The past 20 years have also seen a revolution in the methodology of establishing relationships, and scientists have proposed many new theories regarding the factors controlling the rate and direction of evolution. The useof the fossil record in determining the polarity and ho- mology of characters is emphasized throughout the book as a necessary step in establishing phylogenies. I describe numerous patterns of radiation that can be used to test current hypotheses regarding the processes and con- straints that govern the course of evolution. This book reflects the current direction of paleon- tological research by emphasizing functional anatomy and the interactions between extinct organisms and their en: Vironment. in addition to considering the anatomy and way of life of each of the major vertebrate groups, I have paid special attention to transitions between groups. 1 have also focused on periods of vertebrate history during which the fossil record is sparse and the avaitable evidence has led to conflicting hypotheses of relationship. Thope that this book will not only provide knowledge of our current understanding of the history of vertebrates but will also serve to stimulate further research into the ‘many remaining problems of relationships and evolution- ay processes. ACKNOWLEDGMENTS In a discipline as broad as vertebrate paleontology, ‘one worker can be personally familiar with only a very small portion of the current research, My own studies have focused on a few families of Paleozoic and easly Mesozoic amphibians and reptiles. For information on the remainder of vertebrate phylogeny, | am indebted to a host of friends and colleagues from this and previous generations. I would like to offer special thanks to my graduate students, past and present: Donald Brinkman, Philip Cur- tie, Stephen Godfrey, the late Malcolm Heaton, Robert Holmes, Robert Reisz, Olivier Rieppel, Denis Walsh, and Carl Wellstead. Their work has contributed substantially, to the chapters on early amphibians and reptiles. ‘Many other scientists have provided suggestions and critical comments on sections of the manuscript that deal, with groups with which they are most familiar. In addi- tion, several have allowed me to make use of unpublished manuscripts and illustrations so that this text could reflect the most recent research in this field. To all these col- leagues I express my gratitude. David Bardack, Gerry Case, the late Robert Denison, David Dincley, Beverly Halstead, Karel F. Liem, Richard Lund, Roger Miles, Alex Ritchie, Bobb Schaeffer, Hans-Peter Schultze, Barbara Stahl, and Rainer Zanger! all provided assistance in the fish chapters Donald Baird, Angela and Andrew Milner, E. C. Olson, Alec Panchen, Tim Smithson, Zdenck Spinar, Keith Thomson, and Peter Vavghn have shared generously the results of their research into the ancestry and relationships, of Paleozoic tetrapods, Alan Charig, Sankar Chatterjee, Ned Colbert, Peter Dodson, Richard Fstes, Gene Gaffney, Peter Galton, Carl Gants, Cris Gow, Wann Langston, John, PREFACE MacIntosh, John Ostrom, Dale Russell, and Rupert Wild have provided literature, information from unpublished studics. and many helpful suggestions as to the anatomy and relationships of the various reptile groups. The chap- ter on flight benefited greatly from the assistance of Alan Feduccia, Peter Houde, Storrs Olson, John Ostrom, and Kevin Padian. Mike Archer, Bill Clemens, A. W. Crompton, Rich- ard Fox, Jim Hopson, Nick Hotton, Zofia Kielan-Jawo- rowska, Farish Jenkins, Jason Lilligraven, and Hans-Die- ter Sues provided much help in documenting the origin of mammals and their Mesozoic radiation. Assistance in the preparation of the chapters chronicling the radiation of placental mammals was provided by Mary Dawson, Philip Gingerich, Robert Hoffmann, Leonard Krishtalka, Malcolm McKenna, Nancy Neff, Michael Novacek, Ken Rose, the late George Gaylord Simpson, Jean Sudre, David Webb, and John Wible. Larry Barnes, Daryl Domning, Fd Mitchell, and Clayton Ray helped in establishing re- lationships among the marine mammals. The final chapter on evolution benefited greatly from suggestions offered by Niles Eldredge, Andrew Knoll, and David Webb. 1 am particularly grateful to Hans-Dicser Sues and Denis Walsh for reading the entire manuscript and offering suggestions as to its improvement, Additional reviews of part or all of the manuscript were provided by Leo Laporte, Everett Lindsay, Robert Emry, Milton Hildebrand, Michael Williams, and Lance Grande. ‘The spirit of chis text owes much to the work of Alfred S. Romer, whose outstanding contribution to our understanding of the evolution of Paleozoic amphibians and reptiles and sheer enthusiasm for paleontology serve as examples for all workers in the field. The evolutionary concepts that are elaborated here were developed pri- marily from the writings and teaching of George Gaylord Simpson, who continues to stand as the major figure in the integration of the fossil record and evolutionary theory. ‘The greatest assistance in preparing this text has been provided by Pamela Gaskill, who drafted nearly all the illustrations. Heide Hanson carried the major task of ‘manuscript preparation and the technical aspects of the figures and bibliography. Aruna Ashtakala provided as- sistance in preparing the generic lists. Marie La Ricca and Delise Alison assisted with typing the text and the generic lists. Access to the paleontological literature was greatly facilitated by the help of the staff of Blacker-Wood Library, Eleanor Maclean, Ann Habbick, and Jennifer Adains Production of this book benefited greatly fram the efforts of the staff of W. H. Freeman and Company, no- rably the project editor, Scott Amerman; designer, Lynn Pieroni; art director, Mike Suh and Julia DeRosa, co- ordinator, composition and manufacturing. The senior editor, Jerry Lyons, offered advice at every stage. ‘Anna Di Turi has provided encouragement and sup- port throughout the preparation of this bookFossils and Relationships The history of vertebrates spans more than 500 million years, from their appearance in the Cambrian seas to the present rich and varied fauna. Their history is recorded ina sequence of fossils that documents their skeleral anat- omy, distribution, and evolutionary change. From the fossil record we can determine the inter- relationships of the modern species and crace the origin ofthe skeletal features that characterize each major group. The pattern of vertebrate evolution provides the best available evidence for determining the major processes of evolucion: the origin of new structures, physiological pro- cesses, and adaptive strategies. Fossils document the rate of evolution and may enable us to establish the degree to which environmental, developmental, or other factors in- fluence its direction. ‘This text will review the anatomy, relationships, and, evolutionary patterns of the major groups of vertebrates from their differentiation in the early Paleozoic throughtheir last major radiation in the early Cenozoic. 1 will emphasize the way in which vertebrates have adapted (0 diverse environments and the challenges faced during ma- jor adaptive shifts. Particular attention will be paid to the earlier and previously less well-known forms that are fun damental to understanding the origin of the anatomical and adaptive patterns of the modern groups. Vertebrate evolution is an interdisciplinary subject, involving aspects of geology, comparative anatomy, phys- iology, and evolutionary theory Koowledge of geological processes helps one to un- derstand how vertebrates are preserved as fossils and why animals from different environments have different like- Tihoods of being preserved. Evidence of changes in the patterns of the major land snasses, mountain chains, oceans, and shallow seas helps us 0 explain the distribution of, extinct vertebrates and to reconstruct the climates in which they lived. Knowledge of the skeletal anatomy of modern ver- tebrates is necessary for the study of the bones and teeth, which provide most of our information regarding extinct animals. Direct evidence of the soft anatomy of extinct ‘genera is rarely preserved; nevertheless, an understanding, of the basic pattems and functions of the muscular, cit- culatory, respirators, reproductive, and nervous systems is necessary to appreciate their way of life In addition to knowledge of adult anatomy, an un- derstanding of embryological structures and develop- mental processes will help in interpreting the possible di- rection of evolutionary changes and the origin of new structures, These geological and biological subjects will be covered briefly in Chapters One and Two. The term evolution as it appears in the title covers two distinct concepts. Throughout the text, vertebrate evolution is sreated in the historical sense of changes in structure, function, and adaptation, Evolution may also be considered fram the viewpoint of the underlying ge- netic changes and procestes of selection and population dynamics that explain how these changes occur. The final, chapter will be devoted to a consideration of structural, developmental, and environmental factors that help to explain rhe known patterns and rates of vertebrate evo- lution, The next ewenty chapters will describe vertebrate his- tory as established from the fossil record and through our knowledge of living species. This story is by no means complete or entirely coherent. The early stages of verte: brave history are poorly known, and significant gaps still, separate many major groups. This text will document our current knowledge of each group and discuss the problems, of establishing their relationships and ways of life. Many aspects of vertebrate evolution remain contentious, and, alternative viewpoints will be considered. Technical papers describing fossil vertebrates appear in many journals. Short notes on exceptionally important discoveries appear in Science and Nature. Longer descrip- tive papers are published in Journal of Vertebrate Pa- FOSSILS AND RELATIONSHIPS leontology, Journal of Paleontology, Palaeontology, Pa- lacontographica, Zoological Journal of the Linnean So- ciety, Philosophical Transactions of the Royal Society, and the publications of many universities and public mu- seum, for example, Bulletin of the American Museum, Bulletin of the Museum of Comparative Zoology, Har- vard. All literature through 1983 is cited in the Bibliog- raphy of Fossil Vertebrates, published by the Society of Vertebrate Paleontology, whose membership includes professional vertebrate paleontologists from all countries, The office of this society is currently associated with the Natural History Museum, 900 Exposition Boulevard, Los Angeles, California, 90007. THE NATURE OF THE FOSSIL RECORD Our understanding of the relationships and ways of life of extinct vertebrates depends on knowledge of their geological age and the natureof the deposits in which they were buried. The geological time scale is shown in Figure 1-1, together with an outline of vertebrate phy- logeny. Detailed subdivisions of the geological periods and diagrams of relationships within cach of the verte- brate groups accompany their descriptions. ‘The processes of fossilization limit the type of infor- mation we can gain from extinct organisms. The bodies of most animals are consumed or scattered by predators and scavengers soon after death, and their bones are bro- ken up and decompose. Perhaps no more than one in a million are so quickly buried that they may become fos- silized. The flesh almost invasiably decays, but the bones may be infiltrated by water carrying sediments and soluble minerals that fill up the large cavities and precipitate in smaller channels once occupied by cells and blood vesse's. In most vertebrate fossils, mineral components of the bone retain their integrity, so that histological details and chem- ical composition are little altered even after hundreds of millions of years. A fossil bone is still bone, but it contains a hard and heavy inflling of other minerals as well. Occasionally, entire animals have been enclosed in sediments that hardened to give an impression of the com- plete body surface. Fossils of dinosaurs that became mum mmifed after death have been preserved in this manner. Impressions of other animals have heen preserved in lava flows and amber. Flattened, coalified remains showing body outlines are found in fine-grained sediments that were presumably deposited in an anaerobic environment, free from decomposing organisms. Occasionally, the soft tissues may be preserved by waters bearing pyrite or silica Histological details of muscle fibers and kidney tubules are presecved in fossil sharks from the Upper Devonian Cleveland shale. Footprints are relatively common trace fossils that provide information about the posture, gait, and other aspects of behavior of extinct animals, and they may pro-CHAPTER - ZS g3 i : 65 |= é al 3 i Hi 4 Q —] g 2 2 el 2» ae 26 7 xo ; x0 aoa '§ & not aE it wal pat § 1 of & i sos ay - al uv a} NY a) OY 590 - Fig 1, TEMPORAL DISTRIBUTION OF VERTEBRATE CLASSES AND THEIR PROBABLE RELATIONSHIPS, The geological time scale used in this and other phylogenies is from Harland et al, 1982 vide information about distribution of groups not locally represented by skeletons. Coprolites, or fossilized feces, may reflect the shape of the internal surface of the intes tines (notably the spiral valve of sharks) and the diet of the animal that produced them. In some cases, coprolites contain small bones of animals that are not otherwise found in a given fauna. Although the actual tissue of the nervous system is never fossilized, we can learn about the surface features of the brain from impressions of the inner surface of the 3 cranial bones. In some cases, natural infillings of the cra nial cavities, or endocasts, are preserved and exposed when the overlying bone is eroded or intentionally removed. In relatively large animals, we can remove the matrix filling the skull and make an artificial mold of the inside surface of the bone. We can also study the inside of small and complex skulls through the use of scrial sections. Endocasts of mammals are especially informative and can provide a close approximation of the original shape and volume of the brain. In contrast, the brains of fish, amphibians, and reptiles do not fully occupy the cranial cavity, and neither the surface features nor the volume can be accurately reconstructed on the basis of the sur rounding bones (Jerison, 1973). Behrensmeyer and Hill (1980) consider other aspects of the relationships between fossils and the sediments in which they are buried in “Fossils in the Making.” Rixon (1976) and the journal Curator discuss collection and preparation of fossil vertebrates. Not all animals have an equal chance of fossilization, which results in significant biases in the fossil record. Generally, animals that live in or near the water have a much greater chance of being quickly buried and pre- served than animals that live in strictly terrestrial envi- ronments, On the other hand, fossils of animals that lived in the depths of the ocean are not usually found, because this part of the earth’s crust is rarely exposed on the land surface, The shallower waters of continental margins and inland seas, in contrast, are ideal areas forthe preservation and subsequent exposure of fossils. The fauna may be rich and the flow of sediments from the land copious. These areas are frequently subject to tectonic activity that can thrust them up into mountains, where sediments erode to expose the enclosed fossils. The fossil record is also rich in deltaic and lagoonal environments closer to shore, but it may be difficult to determine whether the animals actually lived in this area, inhabited a more truly marine environment, or washed in from freshwater streams or lakes, We also find large numbers of animals living at the margin of the sea—in the zone of the surf and tide—but their remains tend to be fragmented by the action of the sea. Fossils of animals living in fresh water are rarer sim: ply because throughout geological history, areas of fresh water have been less extensive and served as basins of deposition for shorter periods of time. Nevertheless, large lake systems have yielded rich faunas, which may record very short time intervals over wide areas. Many localities record the fauna of coal swamps dur- ing the Upper Carboniferous, However, the area occupied by the productive localities is infinitesimal compared with the millions of square miles of coal deposits. Hook and Ferm (1985) provide a useful model for the special con ditions that were responsible for the preservation of ver- tebrates in coal swamps. They attribute their concentra: tion at Linton, Ohio, to reducing anaerobic conditions associated with deep abandoned river channels that weredominated by spores and degraded plant remains. For most of the Mesozoic, the expected fauna of coal swamps is missing from the fossil record. Large river systems, especially where they form deltas on the margins of large lakes or oceans, provide an ex cellent source of fossils from a spectrum of aquatic and semiaquatic environments. The famous Lower Permian redbeds of north central Texas are an example of such a deposit. Low-lying flood plains beyond the banks of large river channels preserve some of the richest fossil accu- mulations. Deposits from this environment in southern Africa range in age from the Upper Permian throughout the Triassic; included is a magnificent fauna of mammal- like reptiles that documents the transition between reptiles, and mammals. The vast assemblage of Cenozoic mam- mals from western North America was preserved under similas conditions. Further from major areas of sedimentation the fossil record is drastically reduced: Small bodies of water, ponds, and streams are only temporary features in the geological time scale. Above the basins of deposition, they are likely to be eroded, rather than being preserved in the sedimen- tary record. Under exceptional conditions such higher land may be preserved as the result of major faults, where large areas are suddenly dropped hundreds or thousands of fect and become basins of deposition. The peculiar fauna of the Upper Permian of Madagascar appears to be preserved in this manner (Currie, 1981). in more unusual but locally rich sources of fossils have been found in cave and fissure fillings (Robinson, 1962) and within the upright tree stumps of the lycopod genus Sigillaria (Carroll et al., 1972). Such modes of pres- ervation tend to accumulate animals not ordinarily found in aquatic deposits and partially correct the bias against fully terrestrial vertebrates. The discovery of fossils depends not only on their initial burial and preservation but also on present-day conditions of exposure and erosion. Deserts and semi deserts provide excellent areas for the discovery of fossils, since they are exposed but not immediately weathered away or covered by vegetation. In contrast, the wet tropics, are among the least profitable areas for collecting, since rapid chemical weathering destroys the bones before they are ever exposed and what remains is quickly covered with vegetation. ‘We know little about the fossil history of large areas of the tropics. Fossils have been collected at the margins of the Arctic and in Antarctica, but large areas remain obscured by ice. ‘The changing positions of the continents and areas of tectonic activity have also led to varying distributions of depositional environments. In addition, some deposits have been completely removed by subsequent erosion, and others are so deeply buried that their fauna may never be exposed. Thus, while some geological periods are repre sented by a wide spectrum of different environments, oth FOSSILS AND RELATIONSHIPS ers have a more limited record. There are very few non- marine deposits of the Lower Carboniferous or the upper part of the Lower Jurassic anywhere in the Northern Hemisphere. Consequently, the knowledge of semia- quatic, amphibious, and terrestrial vertebrates of these ages is extremely limited. In the Paleozoic, the fossil record remains a patchwork—certain geological horizons, geo- graphical areas, and environments of deposition are well represented, while others are completely unknown. The hazards of preservation and subsequent expo- sure impose another bias—against groups of animals that were rare or geographically restricted. This bias is par ticularly unfortunate, since most major evolutionary changes probably occurred in small, isolated populations that were subject to stringent selection pressure (Dob- zhansky et al., 1977; Mayr, 1963; Simpson, 1953). Where information regarding transitional forms is most eagerly sought, it is least likely to be available. We have no in- termediate fossils between rhipidistian fish and early am phibians or between primitive insectivores and bats: oniy a single species, Archaeopteryx lithographica vepresents the transition between dinosaurs and birds. On the other hand, certain genera of fish, amphibians, and reptiles are known from thousands upon thousands of fossils from every continent. EVOLUTION Although fossil vertebrates were very incompletely known in the nineteenth century, remains of dino- saurs, giant marine reptiles, and mammals without mod- em descendants provided Darwin and other biologists with irrefutable evidence of extinction and, less directly, of the process of evolution Evolution might have been accepted without fossil evidence, but fossils now seem inexplicable without evo- lution. Yet the foremost vertebrate paleontologists of the nineteenth century—Cavier, Owen, and Agassiz—did not accept evolution as put forward by Darwin but argued for a succession of creations and extinctions. Until the 1940s, vertebrate paleontologists remained outside the mainstream of evolutionary thought. Simpson (1944) and Jepsen (Jepsen, Mayr, and Simpson, 1949) made the frst important contributions to the modern evolutionary syn- thesis (Mayr and Provine, 1980). Pethaps we should not be surprised that vertebrate paleontologists did not support the prevailing view of slow, progressive evolution but tended to elaborate the- ories involving saltation, orthogenesis, or other vitalistie hypotheses. Most of the evidence provided by the fossil record does not support a strictly gradualistic interpre- tation, as pointed out by Eldredge and Gould (1972), Gould and Eldredge (1977), Gould (1985), and Stanley (1979, 1982). Few contemporary paleontologists would deny that natural selection controls the direction of evolution, butCHAPTER T many would seek additional factors to account for the rapid evolution that characterizes the early diversification and radiation of groups and the early stages in the elab- oration of major new structures. The great longevity of many groups and the minor evolutionary changes they exhibited pose another problem. I will discuss these and ‘other general aspects of vertebrate evolution in the final chapter ESTABLISHING RELATIONSHIPS ‘One of the most challenging aspects of vertebrate paleontology is establishing relationships—between extinct genera and possible living counterparts and among torally extinct groups. Taxonomists since the time of the ancient Greeks have based relationships and systems of classification primarily on overall similarities. Linnaeus and other early taxonomists thought these similarities re- sulted from the orderly creation of life by God. Yet, these similarities so soundly reflected basic, inherited attributes of the organisms that the hierarchical system of classifi- cation developed by Linnaeus could be accepted, with little change, by evolutionary biologists. Until recently, few paleontologists were seriously concerned with the methods of classifications neverthe- less, they developed a workable and relatively consistent system that integrates well with the classification of living vertebrates. HENNIGIAN SYSTEMATICS. During the past 20 years, the writings of Willi Hennig (1965, 1966, 1981) have aroused increasing interest in the methodology of determining relationships and their expression in systems of classification. Hennig sought a more objective method of establishing relationships and 2 pattern of classification that was noe prey to the sub: jective approach of individual taxonomists. He also sought to make classification reflect phylogenetic patterns as closely as possible. This system is referred to as phylogenetic systematics, cladism, or Hemnigian systematics. Wiley (1981) provides the most effective recent review of this method. Paleontologists, including Eldredge and Cracraft (1980), Gaffney (1979), and Patterson (1981), have been among the most active supporters of phylogenetic system atics. Despite the difficulty of applying some of the con- ventions of Hennigian classification to extinet groups, the methods of determining relationships provide an objective basis thar greatly assists in establishing reliable phylogen- The most important aspect of Hennigian method- ology is the emphasis on specialized or derived characters in establishing relationships, in contrast with the use of general similarities, as has been the common practice for ‘many taxonomists. Two groups may share a large number of attributes in common, but only those that are special: ized relative to other more distantly related groups dem- onstrate close relationships. For example, all fish share a great number of features, but sharks and chimaeras ‘hol: ocephalians or ratfish) are unique in having claspers end prismatic calcification of their cartilage, which suggests that sharks and chimaeras are more closely related t0 one another than either are to other fishes. Hennig coined several terms to emphasize the unique aspects of this approach. The term apomorphy refers to a derived or specialized trait; plesiomorphy refers to a primitive trait. An autapomorphy is a specialization unique to one group, and a synapomorphy is a specialized trait shared by two or more groups. A symplesiomorphy is a shared primitive trait. All these terms must be defined in relation to a particular taxonomic level. The presence of a hemipenes (a paired, eversible penis) is an autapomor- phy of the Squamata, the single group that includes lizards and snakes. [cis a synapomorphy of the Lacercilia (lizards) and the Ophidia (snakes). Within cither of these groups, the presence of the hemipenes is a primitive character or a symplesiomorphy. ‘An autapomorphy has no value in establishing re- lationships with other groups that lack this character; the presence of a hemipenes does not contribute to our un- derstanding of the specific relationship between the Squa- ‘mata and other groups of reptiles that lack this trait. Nor is the presence of this structure valuable in establishing relationships among the various lizard or snake groups, all of which possess it ‘The term character may be applied to any recogniz~ able attribute of an organism. The total number of char- acters of any organism may be nearly infinite, but when classifying, we need to consider only the characters that vary among the taxa being studied, The term character state refers to the presence or absence of a particular attribute or to a series of alternative ways in which a trait may be expressed. For example, teeth may be present or absent, or their attachment may be to the side of the jaw (pleurodont), to the edge of the jaw (acrodont), or in sockets (thecodont). The term morphocline refers to char- acters that vary quantitatively within a group, for in stance, total body size or proportionate length of limbs. Hennig applied the term sister groups to two groups united by the presence of one or more synapomorphic characters. Hennig argued that a system of classification. could be based entirely on such pairs of taxonomic groups (Figure 1-2). This depends on the assumption that both Of the sister groups will inherit one or more uniquely derived characters from their immediate ancestors. (How- ever, evolution may not always occur according to this pattern, A conservative lineage might give rise to a series of daughter species as a result of geographical separation of small portions of the population. In this case, divergent6 Figure 1-2. FOUR ALTERNATIVE AND MUTUALLY EXCLUSIVE HYPOTHESES OF PHYLOGENETIC RELATIONSHIP. AMONG THREE TAXA. In (a} a b, and c belang to a monophyletic group if they all express some derived characters (1), aot shared with other 233, Taxa b and e are considered eo be more closely related to each other species and their descendants will not necessarily share any unique derived features with the conservative lineage, Their affinities can then be judged only on the basis of overall similarities.) ‘The term sister group can apply to any taxonomic level. Most taxonomists agree that the origin of major ‘groups, like birds or mammals, can ultimately be traced to a particular speciation event. Thus, we may initially apply the concept of sister groups at the level of the spe cies. Higher taxenomic categories, for example, placental and marsupial mammals, may also be considered as sister groups. In other cases, a single species might be identified asthe sister group ofa larger taxonomic assemblage such as a family or an order. MONOPHYLY A major objective of classification is to unite species shar- ing a common ancestry. Sucl groups are termed mono- phyletic. It would be desirable if we could demonstrate thatall taxonomic groups evolsed from individual species, bur this demonstration is rarely if ever possible. Mono- phyletic groups can be identified by the presence of unique, derived characters. All mammals, for instance, can be recognized by the presence of hair, mammary glands, and three ear ossicles, which are not present in any other vertebrates. Groups are sometimes recognized that are later found to have evolved from two or more distinct ancestors. Such groups are termed polyphyletic. The suborder Pinnipedia was long used to refer to both seals (Phocoidea) and the sea lions and walruses (Otarioidea). We now recognize that each group evolved independently from separate fam: ilies of terrestrial carnivores, rather than having a single common ancestor. Taxonomists try to avoid naming polyphyletic groups, but they cannot always differentiate characters that are uniquely evolved from those that have arisen convergently in two or more groups. FOSSILS AND RELATIONSHI'S than ether age ca aif band © share other derived characters (2) not found in a. Taxon a is considered to be the sister groep of b and Taxa hand care sister groups a eachother. (d) Anuntesoled tricotomy in which no unique characters are shared by any two taxa to the ex clusion of the thd HOMOLOGY AND CONVERGENCE In order for characters to be useful in establishing mon- ophyletic groups and relationships between groups, we must first demonstrate that the characters are truly com parable. Long before the acceptance of evolutionary the- ory, scientists recognized that particular elements of the skeleton of all vertebrates are fundamentally similar to one another. This similarity is clearly evident among ter- restrial vertebrates. The bones of the forelimb are readily recognizable whether the limb is used for walking (as in mammalian carnivores), grasping (as in primates), flying (as in bats), or swimming (as in secondarily aquatic forms such as whales). Homology refers to the fundamental sim- ilarity of individual structures that belong to different species within a monophyletic group. ‘Their homologous nature originates from a common ancestral pattern ‘The homology of a structure, such as the humerus, can be demonstrated on the basis of several criteria: 1, Position—The configuration of the humerus may change significantly, but its position relative to other structures (the pectoral girdle and the lower limb) remains constant. 2. Development—The embryological tissue from which the humerus develops is the same in all tetrapods, and the pattern of its ontogeny is sim- ilar in all groups in which the bone occurs. 3. Evolution—The change in shape of the humerus as revealed by the fossil record can be traced from ‘one group to another with no significant breaks, and the most primitive condition observed in sis ter groups is the most similar. Not all homologous characters have such a similar structure and function. Some homologies would be almost impossible to recognize without information from either developmental patterns or the fossil record. One of the most striking cases is provided by the mammalian earod CHAPTER T ossicles (see Chapter 17). Only a single car ossicle, the stapes, is present in reptiles, but three bones are present inmammals. The two additional bones—the malleus and incus—are homologous with bones that form the jaw articulation in reptiles—the articular and quadrare. This homology was first established as a result of observed changes in position and function of these bones in the embryos of modern mammals and, more recently, was demonstrated in a sequence of fossils from the reptilian- ssammalian transition, Not only are some characters that are very different in structure and function homologous, but other char- acters that appear very similar may be nonhomologous. For example, a structurally similar patella, or knee cap, is present in birds, mammals, and some lizards, yet it was not present in the ancestors of any of these groups and certainly evolved separately in each. Characters that are similar in structure and function but have arisen separately rather than from a common ancestor are termed convergent. Convergence is common ingroups that become adapted to a similar habitat or way of life; the similar body shape of whales, ichthyosaurs, and mosasaurs—all of which had different terrestrial ancestors—resulted from secondary adaptation to an aquatic way of life. ‘A second evolutionary pattern that results in the de- velopment of nonhomologous characters, parallelism, és frequently recognized. Simpson (196 1) distinguished these patterns as follows: Convergence is the development of similar characters separately in two or more lineages with- ‘out a common ancestry pertinent to the similarity. Par: “ allelism is the development of similar characters sepa- rately in two or more lineages of common ancestry on the basis of, or channeled by, characteristics of that an- cestry These wo terms may be useful in characterizing evolutionary patterns, but in practice they are difficult to distinguish, When we attempt to determine specific re lationships between groups, parallelism may be consid- ered a special case within the broader category of con- vergence. Homoplasy is frequently used as a synonym for convergence, A major problem in establishing relation- ships is distinguishing characters that are homologous fom those that have arisen as a result of convergence. ‘We can establish directly whether or not a character ob- served in two groups is homologous only through know!- edge of the groups’ immediate common ancestor. If this form can be recognized and has this character, it is ho- ‘mologous. If the ancestor can he recognized on the basis of other criteria but lacks this character, we may assume itevolved convergently. [Fa common ancestor is not known, convergence might be detected from knowledge of the immediate ancestors of cither group, but homology can- not be demonstrated in this manner. The fossil record is the final arbitrator in determining homology for traits that are capable of being preserved, but this excludes almost all aspects of soft anatomy and physiology. Even among groups with a good fossil record, ancestors are frequently not known, and might not be recognized without knowledge from other sources re- garding the possible homology of structures. Several other lines of evidence may be used to dis- tinguish homologous characters from features that have arisen convergently, All are based on probabilities. If structures or processes are complex and similar, itis sta- tistically unlikely thae they would have evolved as a result of convergence. The two major groups of vertebrate, the Agnatha (or jawless vertebrates) and the jawed verte- brates, have basically similar pzecerns of the brain, cranial nerves, and associated sense organs. Even without direct knowledge of the common ancestor of all vertebrates, we say safely assume that it had a comparable pattern. Sim- ilarly, complex developmental changes are unlikely to be matched exactly in groups with different ancestral pat terns. ‘A common, rather than multiple, origin of a trait may also be considered more likely if many related forms exhibit the same condition, For example, all placeacal mammals have a fundamentally similar pazeern of repro- duction, involving, a complex system of endocrine control and many similar features of the placenta, Since these ‘occur consistently in all modern mammalian orders, it seems much more likely that this pattern developed in a ‘common ancestral stock, rather than evolving separately in each group. Similar lines of reasoning can be used to support the homology of elements known in fossil groups where spe- cific ancestry has not been established. Pacterson (1982) advanced an extension of these arguments, suggesting that Any relationships established on the basis of orher criteria may be used to determine the homologous nature of par- ticular characters. If monophyletic groups reflect common ancestry, then any characters they exhibit are likely to be homologous. This argument is based only on probability and is much les significant in establishing homology than evidence from the fossil record, because it provides no specific information regarding individual characters. PARSIMONY. All means of establishing homology ocher than those in- volving developmental patterns and the fossil record are based on the principle of parsimony—that itis more log- ical to accept a hypothesis that depends on a small number Of processes rather than one based on a large number of independent processes. Cladists have applied this princi- ple broadly 0 judge alternative relationships on the basis of the relative number of derived characters that they share. In general, this principle is logical and to some degree underlies all scientific thinking. However, we may question the degree to which it is applicable to esrsbiishing phylogenies. In the case of phylogenetic analysis, the use of parsimony is based on the assumption that most char-acters evolved only once and that convergence is rare Surprisingly, supporters of this doctrine have never tested this assumption. In contrast, biologists working with both modern and extinct groups argue that convergence is very common (Cain, 1982; Carroll 1982). Arguments for the close re- lationship of groups based only on the common presence of derived featares are of little value, if convergence is equally or more common than the unique origin of derived characters. ‘There is no simple correlation between elapsed rime and the amount of evolutionary change, but in general, more changes are likely to have taken place in groups that share only a very distant common ancestry than in those that diverged froma common ancestry more recently. ‘The number of features resulting from convergence will almost inevitably increase during evolutionary history, while the number of characters that they share as a result of their common ancestry cannot inerease, and may decrease, if synapomorphies are lost or altered. Hence, arguments based on parsimony are increasingly less likely to be cor- reet the more distant the proposed common ancestor is in time, ‘Arcempts to determine the relationships of groups on the basis of characters exhibited only in their modern representatives may result in phylogenies that are far dif- ferent from those established on the basis of the fossil record. Two striking examples are provided by the recent works of Gardiner (1982) and Rosen et al. (1981). When attempting to establish relationships of any group with a fossil record, we must emphasize the earliest known members, because they have had the shortest amount of time to evolve new characters since their initial diver- gence, Hence, they should provide us with the best op- portunity to idemtfy the derived features that they share with their closest sister group. Characters that evolve within a group, rather than being present in its most primitive members, have no value in establishing its relationship with a possible sister group. POLARITY If we emphasize the use of derived, rather than prim- itive, characters in establishing relationships, we must devise a means for determining the direction of evolu: tionary change of those characters. Within a series of related species, the number of teeth may differ widely. Is the number of teeth increasing or decreasing? Is a small ‘or a large number of teeth likely to be the peimitive con- dition? The direction of evolutionary change is referred to as polarity. Determining the direction of the polarity of character states or morphoclines is very important to all evolution- FOSSILS AND RELATIONSHIPS. ary biologists, and many authors have written on this subject in recent years (Kluge, 1977s Patterson, 1981; Wiley, 1981) STRATIGRAPHIC SEQUENCE The most obvious way a paleontologist can determine polarity is by the order in which characters appear in the fossil record. If the fossil record is relatively complete or at least representative of the actual pattern of evolution within a group, the direction of evolutionary change should be evident from the sequence of fossils. Paleozoic sharks are in general more primitive than the living shark groups and among mammals, the character states exhibited by Paleocene genera are consistently more primitive than those of animals from the later Tertiary. Butler (1982) recently commented on the question of polarity as applied to the configuration of mammalian molars: Cope (1883) found that in the oldest Tertiary mam- malian fauna, from the Lower Palacocene, .... 38 ‘our of 41 species had upper molars with three prin: cipal cusps, and 35 of these were ériangular, whereas many Focene and later mammals had quadeate mo- lars with four cusps. In this way he was able 0 recognize the tritubercular type as the ancestral form. ‘of upper molar. On the other hand, paleontologists have noted the difficulty of relying exclusively on the fossil record to determine the sequence of anatomical changes (Schaeffer, Hecht, and Ekiredge, 1972), and cladists in general have been suspicious of the fossil record and minimize its im- portance in establishing polarity (Hennig, 1981; Patter- son, 1981; Wiley, 1981). The degree of completeness of the fossil record differs widely from group to group and from one horian to another in the history of any particular group. Because many species and genera have relatively long time spans, wwe cannot assume that the earliest appearance of a par ticular character state indicates that it is the most prim- itive. However, in groups with an extensive fossil record, itis probable that the most primitive character states will be expressed in the older fossils and for derived characters to be more common in later forms. As the fossil Fecord improves, it will better reflec the actual sequence of evo- lution (Fortey and Jefferies, 1982). Despite the difficulties Of interpreting the fossil record, itis the final arbitrator in establishing the antiquity of groups and the distribution of character states (Paul, 1983). However, determining the direction of polarity from the fossil record requires some degree of prios knowledge of the relationships of the groups being investigated. A particular character or character state may have evolved at different times in different lineages. The early appear- ance of a character in one group does not necessarily“q CHAPTER I indicate that it represents the ancestral condition in an- other group that appears later in the fossil record. The earliest known vertebrates have an extensive external car apace, but this does not necessarily indicate that the pres- ence of a carapace was also primitive for sharklike fish, which appear at a later time in the fossil record. This, problem can be minimized if the monophyletic nature of ‘groups can be established first on the basis of other char: acters. However, in practice most significant taxonomic problems involve assemblages in which both their mon- ophyletic nature and the polarity of important characters are in question. ONTOGENY Since the recognition of the evolutionary relationship of ‘organisms, it has been evident that the process of em- bryological development generally parallels phylogeny. Terrestrial vertebrates go through a stage in which the have fshlike gill slits, and mammals go through stages in which elements of their circulatory and excretory systems resemble those seen in the early development of fish, am- phibians, and reptiles before they achieve the defiitive mammalian pattern. Haeckel (1866) summarized this phenomenon with the expression “Ontogeny recapitu- lates phylogeny.” In some cases, this phenomenon enables us to deter: mine the relationships of an animal on the basis of early growth stages, when the adult is so specialized that its + affinities are not readily recognized. The relationship of tunicates(urochordates) with vertebrates would be almost impossible to guess on the basis of the adult structure, but the larvae exhibit a notochord and dorsal hollow nerve cord that demonstrate their affinities with other chordates. Gould (1977) reviewed the relationships between on togeny and phylogeny, poincing out problems in a literal interpretation of Haeckel’s phrase while discussing other aspects of development that may be significant in under- standing phylogenetic change. Fink (1982) stressed the importance of ontogenetic change as a means of establishing the direction of evo: lution. ‘The study of ontogeny is not limited to living species, because changes of the skeleton during growth are frequently seen in fossils. Among carly amphibians and reptiles, the pattern of fusion of tarsal elements, that canbe followed ontogenetically, closely parallels the same process when viewed phylogenetically and provides con- vincing evidence of the homology of elements in the two lasses. The process of neoteny or paedomorphosis, in which juvenile characters are retained in sexually mature indi- viduals, results in an altered ontogenetic sequence relative to that of related groups, but such changes can usually be recognized by the polarity of other traits. OUTGROUP COMPARISON Cladists generally stress outgroup comparison as one of the most important single means of establishing the po- larity of traits. This method is usually discussed in terms of contemporary, living groups, but itis equally or even more convincingly applied to groups with a fossil record. Within a particular group, characters or character states are judged to be derived if they do not appear in other, closely related groups (the outgroups} and primitive if they do, The presence of hair in mammals is clearly a derived character, since itis not possessed by any other vertebrate class. The presence of marsupial bones in both marsupials and monotremes suggests that this feature may be prim- itive for mammals; the absence of these bones is a spe- cialization of placentals, To establish polarity, the ideal outgroup would be the ancestors of the group being studied. The polarity of character change among early amphibians can be estab- lished on the basis of the anatomy of their putative ances- tors, the chipidistian fish. On the other hand, the specific nature of the interrelationships among the various fossil hominids is difficult to assess because we lack knowledge of their immediate ancestors from the Pliocene and thus have difficulty determining which features of the various australopithecine species are likely to be primitive or 5

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