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  • 1.* * * * * *Wonderful LifeThe Burgess Shale and the Nature of HistorySTEPHEN JAY GOULDW. W. NORTON & COMPANY. NEW YORK LONDONCopyright (c) 1989 by Stephen Jay GouldAll rights reservedPrinted in the United States of America.First published as a Norton paperback 1990The text of this book is composed in 10/2/13 Avanta, with display type set inFenice 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 FrostBallantine.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 DataGould, Stephen Jay.Wonderful life: the Burgess Shale and the nature of history / Stephen JayGould.p. cm. Bibliography: p. Includes index.1. Evolution-History. 2. Invertebrates, Fossil. 3. Paleontology-Cambrian. 4.Paleontology-British Columbia-Yoho National Park. 5. Burgess Shale. 6.Paleontology-Philosophy. 7. Contingency (Philosophy) 8. Yoho National Park(B.C.) I. Title.QE770.G67 1989560.9-dc19 88-37469ISBN 0-393-30700-XW.W. Norton & Company, Inc., 500 Fifth Avenue, New York, N. Y 10110W.W. Norton & Company Ltd., 10 Coptic Street, London WC1A 1PU11 12 13 14 15 16 17 18 19 20TO NORMAN D. NEWELLWho was, and is, in the most nobleword of all human speech, my teacherContentsPreface and AcknowledgmentsPage 2

2. CHAPTER I. The Iconography of an Expectation--a PROLOGUE IN PICTURES---the LADDER AND THE CONE: ICONOGRAPHIES OF PROGRESS-REPLAYING LIFES TAPE: THE CRUCIAL EXPERIMENT-_Inset_: The Meanings of Diversity and DisparityCHAPTER II. A Background for the Burgess Shale-LIFE BEFORE THE BURGESS: THE CAMBRIAN EXPLOSION AND THE ORIGIN OF ANIMALS-LIFE AFTER THE BURGESS: soft-bodied FAUNAS AS WINDOWS INTO THE PAST---the SETTING OF THE BURGESS SHALE--WHERE--WHY: THE MEANS OF PRESERVATION--WHO, WHEN: THE HISTORY OF DISCOVERYCHAPTER III. Reconstruction of the Burgess Shale: Toward a New View of Life--a QUIET REVOLUTION--a METHODOLOGY OF RESEARCH---the CHRONOLOGY OF A TRANSFORMATION-_Inset_: Taxonomy and the Status of Phyla-_Inset_: The Classification and Anatomy of ArthropodsThe Burgess Drama-Act 1. _Marrella_ and _Yohoia_: The Dawning and Consolidation of Suspicion,1971-1974---the Conceptual World That Whittington Faced--_Marrella_: First Doubts--_Yohoia_: A Suspicion Grows-Act 2. A New View Takes Hold: Homage to _Opabinia_, 1975-Act 3. The Revision Expands: The Success of a Research Team, 1975-1978--Setting a Strategy for a Generalization--Mentors and Students--Conway Morriss Field Season in Walcotts Cabinets: A Hint Becomes aGenerality, and the Transformation Solidifies--Derek Briggs and Bivalved Arthropods: The Not-So-Flashy butJust-As-Necessary Final Piece-Act 4. Completion and Codification of an Argument: _Naraoia_ and _Aysheaia_,1977-1978-Act 5. The Maturation of a Research Program: Life after _Aysheaia_,1979-Doomsday (There Are No Final Answers)---the Ongoing Saga of Burgess Arthropods---Orphans and Specialists----a Present from Santa Claws--Continuing the March of Weird Wonders---_Wiwaxia_---_Anomalocaris_-Coda-SUMMARY STATEMENT ON THE BESTIARY OF THE BURGESS SHALE--DISPARITY FOLLOWED BY DECIMATION: A GENERAL STATEMENT--ASSESSMENT OF GENEALOGICAL RELATIONSHIPS FOR BURGESS ORGANISMSPage 3 3. ---the BURGESS SHALE AS A CAMBRIAN GENERALITY--PREDATORS AND PREY: THE FUNCTIONAL WORLD OF BURGESS ARTHROPODS---the ECOLOGY OF THE BURGESS FAUNA---the BURGESS AS AN EARLY WORLD-WIDE FAUNA---the TWO GREAT PROBLEMS OF THE BURGESS SHALE---the ORIGIN OF THE BURGESS FAUNA---the DECIMATION OF THE BURGESS FAUNACHAPTER IV. Walcotts Vision and the Nature of History---the BASIS FOR WALCOTTS ALLEGIANCE TO THE CONE OF DIVERSITY---a BIOGRAPHICAL NOTE---the MUNDANE REASON FOR WALCOTTS FAILURE---the DEEPER RATIONALE FOR WALCOTTS SHOEHORN---WALCOTTS PERSONA---WALCOTTS GENERAL VIEW OF LIFES HISTORY AND EVOLUTION-----the BURGESS SHOEHORN AND WALCOTTS STRUGGLE WITH THE CAMBRIAN EXPLOSION---the BURGESS SHALE AND THE NATURE OF HISTORY-_Inset_: A Plea for the High Status of Natural HistoryCHAPTER V. Possible Worlds: The Power of "Just History"--a STORY OF ALTERNATIVES-GENERAL PATTERNS THAT ILLUSTRATE CONTINGENCY---the BURGESS PATTERN OF MAXIMAL INITIAL PROLIFERATION--MASS EXTINCTION--SEVEN POSSIBLE WORLDS---EVOLUTION OF THE EUKARYOTIC CELL-----the FIRST FAUNA OF MULTICELLULAR ANIMALS-----the FIRST FAUNA OF THE CAMBRIAN EXPLOSION-----the SUBSEQUENT CAMBRIAN ORIGIN OF THE MODERN FAUNA-----the ORIGIN OF TERRESTRIAL VERTEBRATES---PASSING THE TORCH TO MAMMALS-----the ORIGIN OF _Homo sapiens_--AN EPILOGUE ON _PIKAIA_BibliographyCreditsIndex [not scanned]Preface and AcknowledgmentsThis book, to cite some metaphors from my least favorite sport, attempts totackle one of the broadest issues that science can address--the nature ofhistory itself--not by a direct assault upon the center, but by an end runthrough the details of a truly wondrous case study. In so doing, I follow thestrategy of all my general writing. Detail by itself can go no further; at itsbest, presented with a poetry that I cannot muster, it emerges as admirable"nature writing." But frontal attacks upon generalities inevitably lapse intotedium or tendentiousness. The beauty of nature lies in detail; the message,in generality. Optimal appreciation demands both, and I know no better tacticthan the illustration of exciting principles by well-chosen particulars. Page 4 4. My specific topic is the most precious and important of all fossillocalities--the Burgess Shale of British Columbia. The human story ofdiscovery and interpretation, spanning almost eighty years, is wonderful, inthe strong literal sense of that much-abused word. Charles Doolittle Walcott,premier paleontologist and most powerful administrator in American science,found this oldest fauna of exquisitely preserved soft-bodied animals in 1909.But his deeply traditionalist stance virtually forced a conventionalinterpretation that offered no new perspective on lifes history, andtherefore rendered these unique organisms invisible to public notice (thoughthey far surpass dinosaurs in their potential for instruction about lifeshistory). But twenty years of meticulous anatomical description by threeEnglish and Irish paleontologists, who began their work with no inkling of itsradical potential, has not only reversed Walcotts interpretation of theseparticular fossils, but has also confronted our traditional view aboutprogress and predictability in the history of life with the historianschallenge of contingency--the "pageant" of evolution as a staggeringlyimprobable series of events, sensible enough in retrospect and subject torigorous explanation, but utterly unpredictable and quite unrepeatable. Windback the tape of life to the early days of the Burgess Shale; let it playagain from an identical starting point, and the chance becomes vanishinglysmall that anything like human intelligence would grace the replay.But even more wonderful than any human effort or revised interpretation arethe organisms of the Burgess Shale themselves, particularly as newly andproperly reconstructed in their transcendent strangeness: _Opabinia_, with itsfive eyes and frontal "nozzle"; _Anomalocaris_, the largest animal of itstime, a fearsome predator with a circular jaw; _Hallucigenia_, with an anatomyto match its name.The title of this book expresses the duality of our wonder--at the beauty ofthe organisms themselves, and at the new view of life that they have inspired._Opabinia_ and company constituted the strange and wonderful life of a remotepast; they have also imposed the great theme of contingency in history upon ascience uncomfortable with such concepts. This theme is central to the mostmemorable scene in Americas most beloved film--Jimmy Stewarts guardian angelreplaying lifes tape without him, and demonstrating the awesome power ofapparent insignificance in history. Science has dealt poorly with the conceptof contingency, but film and literature have always found it fascinating._Its a Wonderful Life_ is both a symbol and the finest illustration I knowfor the cardinal theme of this book--and I honor Clarence Odbody, GeorgeBailey, and Frank Capra in my title.The story of the reinterpretation of the Burgess fossils, and of the new ideasthat emerged from this work, is complex, involving the collective efforts of alarge cast. But three paleontologists dominate the center stage, for they havedone the great bulk of technical work in anatomical description and taxonomicplacement--Harry Whittington of Cambridge University, the worlds expert ontrilobites, and two men who began as his graduate students and then builtbrilliant careers upon their studies of the Burgess fossils, Derek Briggs andSimon Conway Morris.I struggled for many months over various formats for presenting this work, butfinally decided that only one could provide unity and establish integrity. Ifthe influence of history is so strong in setting the order of life today, thenI must respect its power in the smaller domain of this book.The work of Whittington and colleagues also forms a history, and the primarycriterion of order in the domain of contingency is, and must be, chronology.The reinterpretation of the Burgess Shale is a story, a grand and wonderful Page 5 5. story of the highest intellectual merit--with no one killed, no one eveninjured or scratched, but a new world revealed. What else can I do but tellthis story in proper temporal order? Like _Rashomon_, no two observers orparticipants will ever recount such a complex tale in the same manner, but wecan at least establish a groundwork in chronology. I have come to view thistemporal sequence as an intense drama--and have even permitted myself theconceit of presenting it as a play in five acts, embedded within my thirdchapter.Chapter I lays out, through the unconventional device of iconography, thetraditional attitudes (or thinl