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WONDERFUL LIFE the Burgess Shale and Lhe Nature of Hislory

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WONDERFUL LIFE the Burgess Shale and Lhe Nature of Hislory BY STEPHEN JAY GOULD IN NORTON PAPERBACK EVER SINCE DARWIN Reflections in Natural Hislory THE PANDA'S THUMB More Reflections in Natural History THE MISMEASURE OF MAN HEN'S TEETH AND HORSE'S TOES Furlher ReflectUms in Natural Hislory THE FLAMINGO'S SMILE Reflections in Natural History AN URCHIN IN THE STORM Essays about Books and Ideas ILLUMINATIONS A Besliary (with R. W. Purcell) WONDERFUL LIFE The Burgess Shale and lhe Nature of Hislory BULLY FOR BRONTOSAURUS Reflections in Natural Hislory FINDERS, KEEPERS Treasures and Oddities of Natural Hislory Collectors from Peter the Great to Louis Agassiz (with R. W. Purcell) Wonderful Life w . W . NORTON & COMPANY' NEW YORK· LONDON Wonderful Life The Burgess Shale and the Nature of History STEPHEN JAY GOULD Copyright C 19!!9 by St<.pl<~l Jay Could All rights rt'SCl\l.'<I. Prink-<i in the Unik-<i Slilt~ of America. The text of this boobs composed in 10 Ih/ 13 Avanta, with display type set in Fenice Light. Composition and manufacturing by The Haddon Craftsmen, Inc. Book design by Antonina Krass. "Design" copyright 1936 by Robert Frost and renewed 1964 by Lesley Frost Ballantine. Reprinted from The Poetry of Robert Frost, edited by Edward Connery Lathem, by permission of Henry Holt and Company, Inc. Library of Congress Cataloging· in· Publication Data Could, Stephen Jay. Wonderful life: the Burgess Shale and the nature of history / Stephen Jay Could. p. em. Bibliography: p. Includes index. I. Evolution-History. 2. Invertebrates, Fossil. 3. Paleontology-Cambrian. 4. Paleontology-British CoIumbia-Yoho National Park. 5. Burgess Shale. 6. Paleontology-Philosophy. 7. Contingency (Philosophy) 8. Yoho National Park (B.C.) I. Title. QETIO.C67 1989 560'.9---«19 83-37469 ISHN II-W,.,n7IMI·X W. W. Nonon & Company. Inc. SOO Fifth Avenue:. New York. N.Y. 10110 www.wwnonon.com W. W. Nonon & Company Ltd. Castle ~Iousc. 7Sn6 Wells Street. London WIT 3QT 21 22 23 24 25 26 27 28 29 30 To NORMAN D. NEWELL Who was, and is, in the most noble word of all human speech, my teacher Contents Preface and Acknowledgments I3 CHAPTER I. The Iconography of an Expectation 23 A PROLOGUE IN PICTURES 23 THE UDDER AND THE CONE: ICONOGRAPHIES OF PROGRESS 27 REPLAYING LIFE'S TAPE: THE CRUCIAL EXPERIMENT 45 Inset: The Meanings of Diversity and Disparity 49 CHAPTER II. A Background for the Burgess Shale 53 LIFE BEFORE THE BURGESS: THE CAMBRIAN EXPLOSION AND THE ORICIN OF ANIMALS 53 LIFE AFrER THE BURGESS: SoFr-BoDIED FAUNAS AS WINDOWS INTO THE PAST 60 THE SETTINC OF THE BURGESS SHALE 64 WHERE 64 WHY: THE MEANS OF PRESERVATION 69 WHO, WHEN: THE HISTORY OF DISCOVERY 70 10 I CONTENTS CHAPTER III. Reconstruction of the Burgess Shale: Toward a New View of Life 79 A QUIET REVOLUTION 79 A METHODOLOGY OF RESEARCH 84 THE CHRONOLOGY OF A TRANSFORMATION 97 Inset: Taxonomy and the Status of Phyla 98 Inset: The Classification and Anatomy of Arthropods 102 The Burgess Drama 107 Act I. Manella and Yo/lOia: The Dawning and Consolidation of Suspicion, 1971-1974 107 TIle Conceptual World That Whittington Faced 107 Manella: First Doubts 113 Yohoia: A Suspicion Grows 121 Act 2. A New View Takes Hold: Homage to OpabinUz, 1975 124 Act 3. TIle Revision Expands: The Success of a Research Team, 1975-1978 136 Setting a Strategy for a Generalization 136 Mentors and Students 139 Conway Morris's Field Season in Walcott's Cabinets: A Hint Becomes a Generality, and the Transformation Solidifies 141 Derek Briggs and Bivalved Arthropods: The Not-So-F1ashy but Just-As-Neccssary Final Piece 157 Act 4. Completion and Codification of an Argument: Naraoia and Aysheaia, 1977-1978 164 Act 5. The Maturation of a Research Program: Life after Ayshedia, 1979-Doomsday (TIlere Are No Final Answers) 172 The Ongoing Saga of Burgess Arthropods 173 Orphans and Specialists 173 A Prescnt from Santa Claws 185 Continuing the March of Weird Wonders 188 Wiwaxia 189 Anomalocoris 194 Coda 206 CONTENTS III SUMMARY STATEMENT ON THE BESTIARY OF THE BURCESS SHALE 207 DISPARITY FOLLOWED BY DECIMATION: A GENERAL STATEMENT 207 ASSESSMENT OF GENEALOGICAL RELATIONSHIPS FOR BURGESS ORGANISMS 212 THE BURCESS SHALE AS A CAMBRIAN GENERALITY 218 PREDATORS AND PREY: THE FUNcrlONAL WORLD OF BURGESS ARTHROPODS 219 THE ECOLOGY OF THE BURGESS FAUNA 222 THE BURGESS AS AN EARLY WORLD-WIDE FAUNA 224 THE Two GREAT PROBLEMS OF THE BURCESS SHALE 227 THE ORIGIN OF THE BURGESS FAUNA 228 THE DECIMATION OF THE BURGESS FAUNA 233 CHAPTER IV. Walcott's Vision and the Nature of History 240 THE BASIS FOR WALCOTT'S ALLEGIANCE TO THE CoNE OF DIVERSITY 240 A BIOGRAPHICAL NOTE 240 THE MUNDANE REASON FOR WALCOTT'S FAILURE 243 THE DEEPER RATIONALE FOR WALCOTT'S SHOEHORN 253 WALCOTT'S PERSONA 253 WALCOTT'S GENERAL VIEW OF LIFE'S HISTORY AND EVOLUTION 257 THE BURGESS SHOEHORN AND WALCOTT'S STRUGGLE WITH THE CAMBRIAN EXPLOSION 263 THE BURCESS SHALE AND THE NATURE OF HISTORY 277 Inset: A Plea for the High Status of Natural History 280 CHAPTER V. Possible Worlds: The Power of "Just History" 292 A STORY OF ALTERNATIVES 292 GENERAL PATTERNS THAT ILLUSTRATE CONTINCENCY 299 THE BURGESS PATTERN OF MAXIMAL INITIAL PROLIFERATION 301 MASS EXTINcrlON 305 12 I CONTENTS SEVEN POSSIBLE WORLDS 309 EVOLUTION OF THE EUKARYOTIC CELL 309 THE FIRST FAUNA OF MULTICELLULAR ANIMALS 311 THE FIRST FAUNA OF THE CAMBRIAN EXPLOSION 314 THE SUBSEQUENT CAMBRIAN ORIGIN OF THE MODERN FAUNA 316 THE ORIGIN OF TERRESTRIAL VERTEBRATES 317 PASSING THE TORCH TO MAMMALS 318 THE ORIGIN OF Homo sapiens 319 AN EPILOGUE ON PI K A I A 321 Bibliography 325 Credits 333 Index 337 Preface and Acknowledgments This book, to cite some metaphors from my least favorite sport, attempts to tackle one of the broadest issues that science can address-the nature of history itself-not by a direct assault upon the center, but by an end run through the details of a truly wondrous case study. In so doing, I follow the strategy of all my general writing. Detail by itself can go no further; at its best, presented with a poetry that I cannot muster, it emerges as admirable "nature writing." But frontal attacks upon generali­ ties inevitably lapse into tedium or tendentiousness. The beauty of nature lies in detail; the message, in generality. Optimal appreciation demands both, and I know no better tactic than the illustration of exciting principles by well-chosen particulars. My specific topic is the most precious and important of all fossil local i­ ties-the Burgess Shale of British Columbia. The human story of discovery and interpretation, spanning almost eighty years, is wonderful, in the strong literal sense of that much-abused word. Charles Doolittle Walcott, premier paleontologist and most powerful administrator in American sci­ ence, found this oldest fauna of exquisitely preserved soft-bodied animals in 1909. But his deeply traditionalist stance virtually forced a conventional interpretation that offered no new perspective on life's history, and there­ fore rendered these unique organisms invisible to public notice (though they far surpass dinosaurs in their potential for instruction about life's history). But twenty years of meticulous anatomical description by three 1 3 14 I PREFACE AND ACKNOWLEDGMENTS English and Irish paleontologists, who began their work with no inkling of its radical potential, has not only reversed Walcott's interpretation of these particular fossils, but has also confronted our traditional view about prog­ ress and predictability in the history of life with the historian's challenge of contingency-the "pageant" of evolution as a staggeringly improbable se­ ries of events, sensible enough in retrospect and subject to rigorous expla­ nation, but utterly unpredictable and quite unrepeatable. Wind back the tape of life to the early days of the Burgess Shale; let it play again from an identical starting point, and the chance becomes vanishingly small that anything like human intelligence would grace the replay. But even more wonderful than any human effort or revised interpreta­ tion are the organisms of the Burgess Shale themselves, particularly as newly and properly reconstructed in their transcendent strangeness: Opa­ binia, with its five eyes and frontal "nozzle"; Anomalocaris, the largest animal of its time, a fearsome predator with a circular jaw; Hallucigenia, with an anatomy to match its name. The title of this book expresses the duality of our wonder-at the beauty of the organisms themselves, and at the new view of life that they have inspired. Opabinia and company constituted the strange and wonderful life of a remote past; they have also imposed the great theme of contin­ gency in history upon a science uncomfortable with such concepts. This theme is central to the most memorable scene in America's most beloved film-Jimmy Stewart's guardian angel replaying life's tape without him, and demonstrating the awesome power of apparent insignificance in his­ tory. Science has dealt poorly with the concept of contingency, but film and literature have always found it fascinating. It's a Wonderful Life is both a symbol and the finest illustration I know for the cardinal theme of this book-and I honor Clarence Odbody, George Bailey, and Frank Capra in my title. The story of the reinterpretation of the Burgess fossils, and of the new ideas that emerged from this work, is complex, involving the collective efforts of a large cast. But three paleontologists dominate the center stage, for they have done the great bulk of technical work in anatomical descrip­ tion and taxonomic placement-Harry Whittington of Cambridge Uni­ versity, the world's expert on trilobites, and two men who began as his graduate students and then built brilliant careers upon their studies of the Burgess fossils, Derek Briggs and Simon Conway Morris.
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