CHEATS AND DECEITS 3 GreatClarendonStreet,Oxford,OX26DP, UnitedKingdom OxfordUniversityPressisadepartmentoftheUniversityofOxford. ItfurtherstheUniversity’sobjectiveofexcellenceinresearch,scholarship, andeducationbypublishingworldwide.Oxfordisaregisteredtrademarkof OxfordUniversityPressintheUKandincertainothercountries ©MartinStevens2016 Themoralrightsoftheauthorhavebeenasserted FirstEditionpublishedin2016 Impression:1 Allrightsreserved.Nopartofthispublicationmaybereproduced,storedin aretrievalsystem,ortransmitted,inanyformorbyanymeans,withoutthe priorpermissioninwritingofOxfordUniversityPress,orasexpresslypermitted bylaw,bylicenceorundertermsagreedwiththeappropriatereprographics rightsorganization.Enquiriesconcerningreproductionoutsidethescopeofthe aboveshouldbesenttotheRightsDepartment,OxfordUniversityPress,atthe addressabove Youmustnotcirculatethisworkinanyotherform andyoumustimposethissameconditiononanyacquirer PublishedintheUnitedStatesofAmericabyOxfordUniversityPress 198MadisonAvenue,NewYork,NY10016,UnitedStatesofAmerica BritishLibraryCataloguinginPublicationData Dataavailable LibraryofCongressControlNumber:2015944361 ISBN 978–0–19–870789–9 PrintedinGreatBritainby ClaysLtd,StIvesplc LinkstothirdpartywebsitesareprovidedbyOxfordingoodfaithand forinformationonly.Oxforddisclaimsanyresponsibilityforthematerials containedinanythirdpartywebsitereferencedinthiswork. For Audrey and Samuel PREFACE Most of us will have encountered one form of deception or another in the naturalworld.It’shardnottobecausedeceptioniseverywhere.Itrangesfrom therelativelyfamiliarsightofacaterpillarcamouflagedonatree,totheway that some orchids elaborately mimic the smell and appearance of female insects. But what’s often less well appreciated is the intricacy, extent, and sometimestheextremityofdeceitandmanipulationusedbyanimals,plants, andevenfungi.Thisisperhapspartlybecausedeceptionoftenoccursinmany habitats that we are somewhat less familiar with (such as the deep sea or tropical rainforests), and because it frequently occurs in sensory modalities thatwearelesswellattunedto(suchasultravioletlightorultrasonicsounds). Inshort,themorewelookfordeceptionandtrickeryinnature,themorewe find it. I have spent most of my career working on various aspects of deception, from camouflage in crabs through to mimicry by cuckoos, and the extent that organisms go to in order to manipulate one another still astounds me. In some regards it shouldn’t, because biologists have long appreciatedthat being successful, in evolutionary terms,is about passingon your genes to the next generation, and if that means tricking and exploiting otherstodoso,thensobeit.Itis,however,thesophisticatedadaptationsand intricacy involved in many forms of deception, and the number of forms of deceptionthatexistinnature,thatneverceasestoamazeme. Thisbookisaboutwhatweknowregardingdeceptioninnature(andwhat wedon’t).Itisaboutthetypesofdeceptionthatexist,howtheywork,andthe historical context and significance that deception has had, and continues to have, in understanding evolution and adaptation. It is also very much about the modern scientific work that has sought to investigate and understand vii PREFACE deception across a wide range of organisms. Each chapter discusses one or twomaintypesorfunctionsofdeceptionbroadlycoveringthreemainareas: obtainingfood, avoidingbeing eaten, andreproduction. Withoutwishing to give too much away, Chapter1 focuses on a couple of specific examples (especiallyhowsomecaterpillarstrickantsintolookingafterthem)tointro- ducesomekeyideasandconceptsinthestudyofdeception,andtoillustrate justhowsophisticateditcanbe.Chapter2discusseshowanimalsmimicother species,oraspectsoftheenvironment,inordertostealfoodfromothersand tocapturetheirprey.ThisthemeiscontinuedintoChapter3,whichdiscusses thewaysinwhichmanyspecies(especiallyspiders)usedeceptivecommuni- cationsignalsandstimulitoactivelylurepreytowardsthemthroughavariety ofapproaches.Thebookthenturnstohowanimalsusedeceptioninorderto avoidbecomingamealthemselves,startingwithvarioustypesofcamouflage (Chapter4), followed by how harmless animals mimic other dangerous spe- ciessothatpredatorsavoidthem(Chapter5),andthenhowotherspeciesuse suddenstartledisplaysandotherdeceptivetacticstocausepredatorstofleeor ignorethem(Chapter6).Chapters7and8moveontothewaysthatanimals, plants, and fungi trick and manipulate individuals of the same and different speciesforreproduction.Westartwithhowbirdsandinsectsdupeothersinto rearingtheiryoung,beforemovingontohowindividualsmanipulatepoten- tialpartnersandrivalsinmating.Chapter9thenbringsthingsbacktogether to highlight some of the key areas and concepts of deception, and looks forwardstowhatwestillneedtounderstandanddiscover. There are several processes that are thought by scientists to drive the evolutionofdeceptioninnature.Definingkeyconceptsinasubjectlikethis isimportantifwearetobeclearaboutwheredifferentprocessesoccur,and how the type of deception discussed actually works and evolves. However, whileimportant,thereisariskthatformaldefinitionscanbecomeoverlydry, andIdidnotwanttointerruptthestoryofhowdifferenttypesofdeception worktoomuchbyintroducingsemanticsfromtheoutset.Instead,Ihavetried to find a balance between discussing the varied types and examples of deceptionandhowtheyworkwithoutbeingtooformal,whilestilloutlining what the key concepts are. Beyond the brief description that follows here, viii PREFACE I have generally introduced key terms, including when they may occur in nature,wheretheyarefirstrelevantinthebook,ratherthanadoptingamore formal set of definitions and concepts from the very outset in Chapter1. Of these, perhaps the most familiar type of deception is mimicry. Here, an indi- vidualresemblesanotherspeciesinsomeway,suchthatitcandeceiveothers into considering it as the ‘wrong’ object type. For example, some species of harmless snake match the banding colours of venomous species to avoid predators—something called Batesian mimicry. Conversely, aggressive mimicry occurs when mimicry is used in an antagonistic way. For example, some insects resemble the colour and shape of flowers to avoid being recognized and avoided by their prey, which frequently visit real flowers. Camouflage is also a widespread and intuitive idea (though much more complex and intri- guingthangenerallyrealized).Frequently,camouflageinvolvesmatchingthe appearanceof theenvironment, orsome specific objectin it (such as adead leaf), sothat apredator(or preyitem) fails todetectorrecognizeit.Thereis one other key concept worth highlighting here too: sensory exploitation. This occurswhenanindividualproducesacommunicationsignalthathasevolved tobehighlyeffectiveinstimulatingthesensorysystemofanotheranimal(of thesameoranotherspecies).Indoingso,theindividualmakingthesignalcan elicitagreaterbehaviouralresponsefromitstargetthanmightotherwisehave been the case. For example, some frogs produce mating calls that have evolvedspecificsoundpropertiesthatstronglystimulatethehearingsensitiv- ity of females, and in doing so increase their chances of mating. Sensory exploitation seems to be a common way that deception arises and works innature. Inprinciple,thisbookcouldhavebeenorganizedbyanothersetofideasor subjects, and we should not think of the topic(s) of each chapter and the notionsdiscussedasisolatedfromoneanother.Onthecontrary,Ihavetried throughout to indicate common concepts and theories and attempt to link them together. Nonetheless, the main themes of each chapter seemed to be the most logical way to organize things; being the broad types of deception that exist and what ultimately they achieve. It should also be said that the comparativelengthofcertainchaptersandextentoftheideasdiscusseddoes ix
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