Gian Antonio Danieli Alessandro Minelli Telmo Pievani Editors Stephen J. Gould: The Scientific Legacy Stephen J. Gould: The Scientific Legacy Gian Antonio Danieli Alessandro Minelli • Telmo Pievani Editors Stephen J. Gould: The Scientific Legacy 123 Editors Gian AntonioDanieli Alessandro Minelli IstitutoVeneto diScienze, Lettere edArti Telmo Pievani Venice Universityof Padua andIstituto Venetodi Italy Scienze, Lettere edArti Padua,Venice Italy ISBN 978-88-470-5423-3 ISBN 978-88-470-5424-0 (eBook) DOI 10.1007/978-88-470-5424-0 SpringerMilanHeidelbergNewYorkDordrechtLondon LibraryofCongressControlNumber:2013941492 (cid:2)Springer-VerlagItalia2013 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation,broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionor informationstorageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilar methodology now known or hereafter developed. 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While the advice and information in this book are believed to be true and accurate at the date of publication,neithertheauthorsnortheeditorsnorthepublishercanacceptanylegalresponsibilityfor anyerrorsoromissionsthatmaybemade.Thepublishermakesnowarranty,expressorimplied,with respecttothematerialcontainedherein. Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Preface May 20th 2012 was the tenth anniversary of Stephen Jay Gould’s death. Palaeontologist at the Museum of Comparative Zoology, Harvard University, eminent evolutionary biologist, science writer, science historian and opinion maker, Gould gave us an extended and revised version of the theory of evolu- tion, his ‘‘Darwinian pluralism’’, which is still today an interesting frame to understand the scientific advancements in many evolutionary fields. His antic- ipating insights about the conjunction of evolution and development, the role of ecological and biogeographical factors in ‘‘punctuated’’ speciation, the need for a multi-level interpretation of the units of selection, the interplay between func- tional pressures and internal constraints in processes like exaptations and span- drels are fruitful current lines of experimental research today. Even his pungent and sometimes very radical controversies against the progressive representations of evolution (especially human evolution), biological determinism, pan-selectionist and a gene-centered view of natural history, or the adaptationist ‘‘just-so-stories’’, have left their mark in contemporary biology. Gould’s ‘‘histories of nature’’ were explorations in the ‘‘nature of history,’’ with wider cultural and philosophical implications, like his crucial concept of contin- gency. Thus, after 10 years of new discoveries and unforeseen advances, it is worthytodiscusstheefficacyandlimitsofGould’spluralismasrenovationofthe Darwinian research program. At the historical location of the Istituto Veneto di Scienze, Lettere ed Arti in Venice, the town of Gould’s ‘‘spandrels of San Marco’’, an international panel of scientistsandphilosophers—includingGould’sclosestfriendsandcolleagueslike Niles Eldredge, Elisabeth Lloyd, and(in video) Richard Lewontin—discussed his evolutionaryandanthropologicallegacy,hisideaofscienceasacomplexrational enterprise, evolving itself and immersed in human society, his proposal for a methodology in historical sciences, and his unmistakable style of writing and argumentation, overcoming the boundaries between science, literature, and art. InGould’sproduction,scientificresearchandcommunicationofscienceweretwo fields of inquiry strictly related by the idea that science is a high expression of human curiosity and culture. The International Meeting was held at the Istituto Veneto, with the collabora- tion of University Ca’ Foscari, on May 10–12th 2012. We thank Maria Turchetto v vi Preface and Elena Gagliasso for their helpful participation in the organizing committee. The Venetian meeting has been the basis for the construction of this volume, which is divided into four parts. The first one—with the contributions of Niles Eldredge, Elisabeth Lloyd, and Telmo Pievani—is focused on the general scien- tific legacy of Stephen J. Gould as an evolutionary biologist: the unpublished history of the birth of Punctuated Equilibria; the role of Gould’s criticism against adaptationism;thestructureofhis‘‘Darwinianpluralism’’.Thesecondpart—with the contributions of T. Ryan Gregory, Alessandro Minelli, Gerd Müller, and MarcelloBuiatti—isdedicatedtothediscussionofGould’stheoreticalinnovations seen from the perspective of genomics and developmental biology: the Gouldian idea of genome as a hierarchical system; the debate about the levels of selection and the ‘‘individual’’ units in evolution; his anticipations of some fundamental ‘‘Evo-Devo’’conceptslikedevelopmentalconstraintsandspandrels;hisintuitions about the complexity of genetic coding and differential mutation rates. The third part—withthecontributionsofIanTattersall,GuidoBarbujani,KlausR.Scherer, and Winfried Menninghaus—deals with the important anthropological legacy of Stephen J. Gould: his advocacy of a highly branching phylogeny of hominids, against any progressive idea of cumulative change in human evolution; his bold fight against biological determinism and the alleged genetic foundations of the concept of ‘‘human races,’’ the evolution of emotions, speech, and music in a Gouldian perspective. The fourth part—with the contributions of Andrea CavazziniandAlbertoGualandi—isfocusedonsomeaspectsofGould’slegacyin human sciences, with reference to the conceptual shifts between economics and evolutionary theory, and the possibilities and limits of Gould’s humanism. The richness of Gould’s production and intellectual inheritance cannot be covered by a single collections of essays. Nevertheless, we hope to add another piecetotherichmosaicofstudiesthattheHarvardevolutionistdeserves.Gould’s ‘‘industry’’ is a mine of historical hints, epistemological proposals, scientific insights, and contentious theories. As Richard Lewontin said in his thoughtful opening address by video conference, Gould’s way of exploring evolution was a mix of pure history and theoretical generalizations, aided by extraordinary com- municational skills and a worldwide reputation. He was so brilliant inventing metaphors(suchas‘‘spandrels’’and‘‘PunctuatedEquilibria’’),thathewasableto depict for professionals, and for the general audience at the same time, the wide frame of the ‘‘multiple generating forces of evolution.’’ He was a forerunner. He challenged several orthodoxies, included the ‘‘ultra- Darwinian’’one.Hebecameastraw-manformanyopponents.Stillnow,heisone of the most quoted evolutionists. During these first ten years his proposals and provocations have had a differential survival, but there are no doubts that his pluralismhasstronglyinfluencedthecurrentdebate.StephenJ.Gouldisapresent- day evolutionist. Gian Antonio Danieli Alessandro Minelli Telmo Pievani Contents Part I Evolutionary Theory Stephen J. Gould in the 1960s and 1970s, and the Origin of ‘‘Punctuated Equilibria’’ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Niles Eldredge Stephen J. Gould and Adaptation: San Marco 33 Years Later . . . . . . 21 Elisabeth A. Lloyd Kinds of Pluralism: Stephen J. Gould and the Future of Evolutionary Theory. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 Telmo Pievani Part II Genome and Development Molecules and Macroevolution: A Gouldian View of the Genome. . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 T. Ryan Gregory Individuals, Hierarchies and the Levels of Selection: A Chapter in Stephen J. Gould’s Evolutionary Theory. . . . . . . . . . . . 73 Alessandro Minelli Beyond Spandrels: Stephen J. Gould, EvoDevo, and the Extended Synthesis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85 Gerd B. Müller Biological Complexity and Punctuated Equilibria. . . . . . . . . . . . . . . . 101 Marcello Buiatti vii viii Contents Part III The Anthropological Legacy Stephen J. Gould’s Intellectual Legacy to Anthropology. . . . . . . . . . . 115 Ian Tattersall Mismeasuring Man Thirty Years Later . . . . . . . . . . . . . . . . . . . . . . . 129 Guido Barbujani Affect Bursts as Evolutionary Precursors of Speech and Music. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147 Klaus R. Scherer Darwin’s Theory of Music, Rhetoric and Poetry. . . . . . . . . . . . . . . . . 169 Winfried Menninghaus Part IV Stephen J. Gould and Human Sciences Beyond (and Without) the Invisible Hand. Conceptual Shifts Between Economics and the Theory of Evolution . . . . . . . . . . . 179 Andrea Cavazzini Stephen J. Gould, Between Humanism and Anti-humanism. Neoteny, Exaptation and Human Sciences . . . . . . . . . . . . . . . . . . . . . 187 Alberto Gualandi Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201 Part I Evolutionary Theory Stephen J. Gould in the 1960s and 1970s, and the Origin of ‘‘Punctuated Equilibria’’ Niles Eldredge Abstract SteveGouldarrivedasabeginninggraduatestudentinthe Department ofGeologyat Columbia Universityinthe Fall of1963.He was oneofagroup of enteringstudentsinterestedinpaleontology,biostratigraphy,paleoecologyand,of course, evolution. Though I was still an undergraduate, I was welcomed into the group—and took part in the field trips and special seminars they organized: especially one on paleontology and evolution whose main inspiration was Steve himself.Mostofthesestudentseventuallywentontohavedistinguishedcareersin paleontology and related fields. Steve’s initial—and perhaps always his favorite—professional passion was mor- phology, development and evolution. He astonished everyone that he would devote an entire year away from his doctoral research to write an exploratory review paper on allometry—inspired by his initial work as an undergraduate with JohnWhiteonthemeaningof‘‘b’’inthefamousequationY = bXk.Stevequickly emerged as a model of the ambitious young professional, encouraging us all to developandpublishresearchprojects—andtobeboldandthinkabouttheoretical issues. He once said to me Why wait until we are 60 before we publish on evolutionary theory? And of course he was right; indeed, sadly, he did not live beyond that very age. The genesis of our 1972 paper Punctuated Equilibria: An Alternative to Phy- leticGradualismhasbeenrecountedseveraltimes,bySteveandbymyselfaswell as by others. The definitive version, in my view, is in the newly published book RereadingtheFossilRecord(2012)byhistorianDavidSepkoski.Iwillreviewthe essential details of our joint participation in Tom Schopf’s GSA Symposium and multi-authoredbook,bothentitledModelsinPaleobiology.Thoughthegistofthe concept of punctuated equilibria was developed in my 1971 paper The Allopatric N.Eldredge(&) AmericanMuseumofNaturalHistory,NewYork,USA e-mail:[email protected] G.A.Danielietal.(eds.),StephenJ.Gould:TheScientificLegacy, 3 DOI:10.1007/978-88-470-5424-0_1,(cid:2)Springer-VerlagItalia2013