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 I. The Iconography of an Expectation 23 CHAPTER 23 A PROLOGUE IN PICTURES THE UDDER AND THE CONE: ICONOGRAPHIES OF PROGRESS 27 45 REPLAYING LIFE'S TAPE: THE CRUCIAL EXPERIMENT Inset: The Meanings of Diversity and Disparity 49 II. A Background for the Burgess Shale 53 CHAPTER LIFE BEFORE THE BURGESS: THE CAMBRIAN EXPLOSION AND 53 THE ORICIN OF ANIMALS LIFE AFrER THE BURGESS: SoFr-BoDIED FAUNAS AS WINDOWS INTO THE PAST 60 64 THE SETTINC OF THE BURGESS SHALE 64 WHERE 69 WHY: THE MEANS OF PRESERVATION 70 WHO, WHEN: THE HISTORY OF DISCOVERY I 10 CONTENTS III. Reconstruction of the Burgess Shale: CHAPTER 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, 19 79-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 III CONTENTS SUMMARY STATEMENT ON THE BESTIARY OF THE BURCESS 207 SHALE 207 DISPARITY FOLLOWED BY DECIMATION: A GENERAL STATEMENT ASSESSMENT OF GENEALOGICAL RELATIONSHIPS FOR BURGESS 212 ORGANISMS 218 THE BURCESS SHALE AS A CAMBRIAN GENERALITY PREDATORS AND PREY: THE FUNcrlONAL WORLD OF BURGESS 219 ARTHROPODS 222 THE ECOLOGY OF THE BURGESS FAUNA 224 THE BURGESS AS AN EARLY WORLD-WIDE FAUNA 227 THE Two GREAT PROBLEMS OF THE BURCESS SHALE 228 THE ORIGIN OF THE BURGESS FAUNA 233 THE DECIMATION OF THE BURGESS FAUNA IV. Walcott's Vision and the Nature of History 240 CHAPTER THE BASIS FOR WALCOTT'S ALLEGIANCE TO THE CoNE OF 240 DIVERSITY 240 A BIOGRAPHICAL NOTE 243 THE MUNDANE REASON FOR WALCOTT'S FAILURE 253 THE DEEPER RATIONALE FOR WALCOTT'S SHOEHORN 253 WALCOTT'S PERSONA WALCOTT'S GENERAL VIEW OF LIFE'S HISTORY AND 257 EVOLUTION THE BURGESS SHOEHORN AND WALCOTT'S STRUGGLE WITH 263 THE CAMBRIAN EXPLOSION 277 THE BURCESS SHALE AND THE NATURE OF HISTORY Inset: A Plea for the High Status of Natural History 280 V. Possible Worlds: The Power of "Just History" 292 CHAPTER 292 A STORY OF ALTERNATIVES 299 GENERAL PATTERNS THAT ILLUSTRATE CONTINCENCY 301 THE BURGESS PATTERN OF MAXIMAL INITIAL PROLIFERATION 305 MASS EXTINcrlON