THE r The Beauty, Elegance, and Strangeness of Insect Societies Bert Holldobler and E. O.Wilson WINNERS OF THE PULITZER PRIZE FOR THE ANTS THE SUPERORGANISM BOOKS BY THE AUTHORS ALSO BY BERT HOLLDOBLER AND EDWARD O. WILSON TheAnts(1990); PulitzerPrize,General Nonfiction, 1991 JourneytotheAnts:AStoryofScientificExploration(1994) ALSO BY BERT HOLLDOBLER ExperimentalBehavioralEcologyandSociobiology',withMartinLindauer,editors (1985) HerbivoryofLeafCuttingAnts,withRainerWirth,HubertHerz, RonaldJ.Ryel, andWolfram Beyschlag(2003) ALSO BY EDWARD O. WILSON TheTheoryofIslandBiogeography,withRobertH.MacArthur(1967); newpreface, 2001 APrimerofPopulationBiology, withWilliam H. Bossert(1971) TheInsectSocieties(1971) Sociobiology: TheNewSynthesis(1975); newedition,2000 OnHumanNature(1978); PulitzerPrize,General Nonfiction, 1979 CasteandEcologyintheSocialInsects,withGeorgeEOster(1978) Genes, Mind, andCulture,withCharlesJ.Lumsden(1981) PrometheanFire:ReflectionsontheOriginofMind,withCharlesJ.Lumsden (1983) Biophilia(1984) SuccessandDominanceinEcosystems: TheCaseoftheSocialInsects(1990) TheDiversityofLife(1992) Naturalist(1994); newedition, 2006 InSearchofNature(1996) Consilience: The UnityofKnowledge(1998) BiologicalDiversity: TheOldestHumanHeritage(1999) TheFutureofLife(2002) PheidoleintheNewWorld:AHyperdiverseAntGenus(2003) FromSoSimpleaBeginning: TheFourGreatBooksofDarwin,editedwith introductions(2005) NatureRevealed:SelectedWritings, 1949-2006(2006) TheCreation:AnAppealtoSaveLifeonEarth(2006) k^c ^•-nfrti"1 Frontispiece: Plate 1.Weaverants(Oecophyllasmaragdina)cooperatein arrangingleavesduringtheconstructionoftheirleaftentnests. Copyright©2009byBertHoUdoblerandEdwardO.Wilson Allrightsreserved PrintedintheUnitedStatesofAmerica FirstEdition Forinformationaboutpermissiontoreproduceselectionsfromthisbook,writeto Permissions,W.W.Norton&Company, Inc.,500FifthAvenue,NewYork, NY 10110 Forinformationaboutspecialdiscounts forbulkpurchases,pleasecontact W [email protected] ManufacturingbyRRDonnelley,Willard,OH BookdesignbyAbbateDesign Productionmanager:AnnaOler LibraryofCongressCataloging-in-PublicationData HoUdobler, Bert, 1936- Thesuperorganism:thebeauty,elegance,andstrangenessofinsectsocieties/BertHoUdobler andEdwardO.Wilson;linedrawingsbyMargaretC.Nelson.— 1sted. p. cm. Includesbibliographicalreferencesandindex. ISBN978-0-393-06704-0(hardcover) 1.Insectsocieties.I.Wilson,EdwardO. II.Title. QL496.H65 2009 595.7'1782—dc22 2008038547 W.W.Norton&Company,Inc. 500FifthAvenue,NewYork, N.Y. 10110 www.wwnorton.com W.W.Norton&CompanyLtd. CastleHouse,75/76WellsStreet,LondonWIT3QT 1234567890 FOR MARTIN LINDAUER, ourcolleagueandfriend, whose pioneeringworkandinspirationinexperimental sociobiobgycontributedgreatlytotheconceptionofan insectsocietyasafunctionalsuperorganism Let him who boasts the knowledge of actuallyexistingthings, first tellusofthe nature ofthe ant. -ST. BASIL CONTENTS NOTE TO THE GENERAL READER XVI CHAPTER 1 THE CONSTRUCTION OF A SUPERORGANISM WhyColoniesAreSuperior 5 TheConstructionofSuperorganisms 6 TheLevels ofOrganization 7 EusocialityandtheSuperorganism 8 ABriefHistoryofInsectSociobiology 10 CHAPTER 2 GENETIC SOCIAL EVOLUTION 15 AnAbridged HistoryoftheGeneticTheoryofSocial Evolution 16 Multilevel NaturalSelection 24 The Evolution ofEusociality 29 Crossing theEusocialityThreshold 31 CountervailingForcesofSelection 42 PassingthePointofNoReturn 42 CHAPTER 3 SOCIOGENESIS 51 TheColonyLife Cycle 53 SocialAlgorithms 53 Self-Organization andEmergence 58 PhylogeneticInertiaandDynamicSelection 60
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