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Beasts of Eden The publisher gratefully acknowledges the generous contribution to this book provided by the General Endowment Fund of the University of California Press Associates. Beasts of Eden walking whales, dawn horses, and other enigmas of mammal evolution David Rains Wallace university of california press berkeley los angeles london University of California Press Berkeley and Los Angeles, California University of California Press, Ltd. London, England © 2004 by the Regents of the University of California Library of Congress Cataloging-in-Publication Data Wallace, David Rains, 1945–. Beasts of Eden : walking whales, dawn horses, and other enigmas of mammal evolution / David Rains Wallace. p. cm. Includes bibliographical references and index. isbn 0-520-23731-5 (alk. paper). 1. Mammals—Evolution. 2. Mammals, Fossil. I. Title. ql708.5.w25 2004 599.138—dc22 2003022857 Manufactured in the United States of America 13 12 11 10 09 08 07 06 05 04 10987654 321 Printed on Ecobook 50 containing a minimum 50% post-consumer waste, processed chlorine free. The balance contains virgin pulp, including 25% Forest Stewardship Council Certified for no old growth tree cutting, processed either tcf or ecf. The sheet is acid-free and meets the minimum requirements of ansi/niso z39.48-1992 (r 1997) (Permanence of Paper). ᭺ϱ to steve fisher, who brings art to science Actually most scientific problems are far better understood by studying their history than their logic. ernst mayr quoted in Leo F. Laporte, George Gaylord Simpson: Paleontologist and Evolutionist contents list of illustrations /xi acknowledgments / xiii prologue. the fresco and the fossil /xv 1 / Pachyderms in the Catacombs /1 2 / Dr. Jekyll and the Stonesfield Jaws /14 3 / The Origin of Mammals /25 4 / The Noblest Conquest /41 5 / Terrible Horns and Heavy Feet /54 6 / Mr. Megatherium versus Professor Mylodon /70 7 / Fire Beasts of the Antipodes /79 8 / Titans on Parade /91 9 / Five-Toed Horses and Missing Links / 104 10 / The Invisible Dawn Man / 115 11 / A Bonaparte of Beasts / 123 12 / Love and Theory / 135 13 / Simpson’s Cynodont-to-Smilodon Synthesis / 145 14 / Shifting Ground / 157 15 / Dissolving Ancestries / 166 16 / Exploding Faunas / 176 17 / The Revenge of the Shell Hunters / 188 18 / Simpson Redivivus / 198 19 / Winds Thieves of the Kyzylkum / 207 20 / The Serpent’s Offering / 216 21 / Anthropoid Leapfrog / 233 epilogue. cenozoic parks / 249 notes / 261 select bibliography / 297 index / 315 illustrations 1. O. C. Marsh’s Dromocyon skeleton, now named Synoplotherium / xix 2. Charles Knight’s restoration of Cope’s Mesonyx /xx 3. Megatherium and Glyptodon (Pleistocene) from Zallinger’s Age of Mammals /2 4. Cuvier’s restorations of Paris gypsum mammals /9 5. Coryphodon and Oxyaena, with Pelycodus (Eocene) from Zallinger’s Age of Mammals /27 6. Caricature by Frederick Waddy of Richard Owen astride a skeletal Megatherium /37 7. Uintatherium and Eobasileus (Eocene) from Zallinger’s Age of Mammals /55 8. Skeletons of Pantolambda and Coryphodon /67 9. Punch’s 1890 cartoon of O. C. Marsh with a uintathere and a titanothere /75 10. Brontops and Palaeolagus with Archaeotherium (Oligocene) from Zallinger’s Age of Mammals /93 11. Erwin Christman’s restoration of titanothere evolution /99 xi 12. E. R. Fulda’s restoration of the giant Mongolian mesonychid Andrewsarchus / 113 13. Smilodon and Bison (Pleistocene) from Zallinger’s Age of Mammals / 121 14. Aepycamelus and Gomphotherium (Miocene) from Zallinger’s Age of Mammals / 124 15. Diatryma and Mesonyx (Eocene) from Zallinger’s Age of Mammals / 154 16. Eomaia scansoria painting by Mark A. Klingler / 214 17. Serpent in tree (Paleocene) from Zallinger’s Age of Mammals / 217 18. Jacob Wortman with dinosaur bones / 222 19. Simpson with a bush baby / 247 20. Rudolph F. Zallinger with his Age of Mammals / 251 xii list of illustrations acknowledgments A number of individuals and institutions helped me write this book. Back in 1993, before I even got the idea, David Webb and Bruce MacFadden of the Florida State Museum of Natural History talked to me about mammal evolution. Spencer Lucas of the New Mexico Museum of Natural History, Brent Breithaupt of the University of Wyoming Museum of Geology, and Greg McDonald, then of Hagerman Fossil Beds National Monument in Idaho, showed me around fossil sites and answered questions in 1997. Alan Rabinowitz gave me advice about a trip to Thailand in 1998, and Mark Graham provided advice and hospitality when I was in that country. David Wake of the U.C. Berkeley Museum of Vertebrate Zoology ad- vised me about sources. Chris Beard, Mary Dawson, and Zhexi Luo of the Carnegie Museum of Natural History answered questions and provided invaluable reference material when I visited there in 2000. Mary Ann Tur- ner of the Yale Peabody Museum spent most of a morning showing me the Zallinger murals, and she, Joyce Gherlone, and other staff members sub- sequently provided much invaluable information and other help. Andrew Petryn helped me with questions about Zallinger’s artistic techniques. David Archibald of the San Diego State University talked to me at length about a relatively little known subject, and generously provided reference material and constructive criticism. Lowell Dingus of the American Mu- seum of Natural History answered questions, and Matthew Pavlick helped xiii me with illustrations. William A. Clemens of the Museum of Paleontology at U.C. Berkeley spent a morning showing me some of their collection, and Patricia Holroyd of the same institution provided much useful infor- mation. Ruthann Knudson of Agate Fossil Beds National Monument helped me with information and illustrative materials. Joan Burns of Wil- liams College generously provided me with a photo of her father, George Gaylord Simpson. This book would have been impossible without the work of the many authors, living and dead, who’ve written about mammal evolution. I’m particularly grateful to writers on evolutionary history, including Martin J. S. Rudwick, Adrian Desmond, Peter J. Bowler, and Ronald Rainger, among others, for the inspiration their work has provided. Vincent Mor- gan and Spencer Lucas were especially helpful in letting me see pre- publication copies of their bulletins on Walter Granger and the Osborn Fayum expedition, and by answering questions that arose from that. And, of course, I’m grateful for the writings of the paleontologists themselves, from Cuvier to the amazingly prolific Simpson. This book also would have been impossible without the services of the University of California at Berkeley, particularly the Biosciences and Earth Sciences libraries, as well as the Doe, Bancroft, and Anthropology libraries. The Geology Library at the Ohio State University, Columbus, was a valu- able resource as well. Finally, I wish to thank my agent, Sandy Taylor, and my editor at the University of California Press, Blake Edgar, for their indispensable help in transforming this project from an ambitious proposal to a manuscript. For misinterpretations, misquotations, factual errors, or other flaws, of course, I take full responsibility. xiv acknowledgments prologue The Fresco and the Fossil If there is a Sistine Chapel of evolution, it is Yale University’s Peabody Mu- seum. Of course, the Peabody’s neo-Gothic brick edifice is less august than the Vatican’s papal shrine. Past a dim corridor off the museum’s vestibule, however, a painting as breathtaking as the Sistine murals covers the wall of a soaring room with an immense landscape of rosy cliffs, exotic vegetation, and life-sized dinosaurs. This is Rudolph Zallinger’s Age of Reptiles, and, like Michelangelo’s murals, it is a fresco, brushed on plaster day after day over a period of years. Like the Sistine’s biblical vision of human creation and judgment, it shows the beginning and end of a world, that of the great saurians that left their bones in western North America. Comparing a big dinosaur picture to Renaissance painting’s supreme achievement seems presumptuous, to be sure, and Rudolph Zallinger is not considered a titan even of modern art. Born to Siberian refugees in 1919, he attended Yale’s School of Fine Arts on scholarships during the Depres- sion, when it was training illustrators, and taught there after graduation. The Peabody hired him to paint the mural in 1943 at $40 a week, because the director felt its Great Hall “resembled a dismal barren cavern devoid of color.” Zallinger took a crash course in paleontology and finished the job in 1947. Three years later, abstractionists purged illustrators from Yale’s art school, and Zallinger might have spent his career doing ads in his home xv town of Seattle if the Peabody had not appointed him “artist in residence,” a position he held—teaching elsewhere—until his death in 1995. The art historian Vincent Scully was speaking outside the mainstream when he complained in 1990 that Zallinger’s work is not “valued as it ought to be by modern critics.” Most critics aren’t aware it has artistic value. In a way, however, Zallinger had more in common with Michelangelo than do the modernists who have prospered from critical and commercial pa- tronage. Great noncommercial institutions—religion, science—patron- ized both muralists, with similar parsimony, and both murals have become icons, endlessly reproduced in popular media. In one stamp issue alone, the U.S. Postal Service printed six million copies of scenes from the Age of Reptiles. And although the modernist canon excludes Zallinger’s mural, its status is more than popular. Scully