Bats: From Evolution to Conservation PDF
Preview Bats: From Evolution to Conservation
Bats This page intentionally left blank Bats From Evolution to Conservation Second Edition John D. Altringham ProfessorofAnimalEcologyandConservation, UniversityofLeeds DRAWINGS BY Tom McOwat and Lucy Hammond Myotismyotis(drawingbyLucyHammond) 1 3 GreatClarendonStreet,Oxfordox26dp OxfordUniversityPressisadepartmentoftheUniversityofOxford. ItfurtherstheUniversity’sobjectiveofexcellenceinresearch,scholarship, andeducationbypublishingworldwidein Oxford NewYork Auckland CapeTown DaresSalaam HongKong Karachi KualaLumpur Madrid Melbourne MexicoCity Nairobi NewDelhi Shanghai Taipei Toronto Withofficesin Argentina Austria Brazil Chile CzechRepublic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore SouthKorea Switzerland Thailand Turkey Ukraine Vietnam OxfordisaregisteredtrademarkofOxfordUniversityPress intheUKandincertainothercountries PublishedintheUnitedStates byOxfordUniversityPressInc.,NewYork #JohnD.Altringham,2011 Alldrawings#TomMcOwatunlessotherwisecredited. Themoralrightsoftheauthorhavebeenasserted DatabaserightOxfordUniversityPress(maker) Firstpublished1996 Secondeditionpublished2011 Allrightsreserved.Nopartofthispublicationmaybereproduced, storedinaretrievalsystem,ortransmitted,inanyformorbyanymeans, withoutthepriorpermissioninwritingofOxfordUniversityPress, orasexpresslypermittedbylaw,orundertermsagreedwiththeappropriate reprographicsrightsorganization.Enquiriesconcerningreproduction outsidethescopeoftheaboveshouldbesenttotheRightsDepartment, OxfordUniversityPress,attheaddressabove Youmustnotcirculatethisbookinanyotherbindingorcover andyoumustimposethesameconditiononanyacquirer BritishLibraryCataloguinginPublicationData Dataavailable LibraryofCongressCataloginginPublicationData Dataavailable TypesetbySPIPublisherServices,Pondicherry,India PrintedinGreatBritain onacid-freepaperby CPIAntonyRowe,Chippenham,Wiltshire ISBN 978–0–19–920711–4(Hbk.) 978–0–19–920712–1(Pbk.) 10 9 8 7 6 5 4 3 2 1 Table of contents Preface vii Acknowledgements ix Introduction—abiologylessonfromthebats xi 1. Evolutionanddiversity 1 Evolution....................................................................................................................................1 Classificationandthedistributionanddiversityofbats ................................................18 Yinpterochiroptera ...............................................................................................................20 Yangochiroptera.....................................................................................................................26 Adaptiveradiation—whyarethePhyllostomidaesonumerousandsodiverse? ......35 2. Flight 37 Somebasicaerodynamics:aerofoilsandflappingflight ................................................37 Theevolutionofflight .........................................................................................................43 Theadvantagesofflightandthedemandsontheflyer:physiological andbiomechanicalaspectsofflight....................................................................................46 Thewingandwhatitdoesinflight....................................................................................50 Ecologicalaspectsofflight ...................................................................................................51 3. Echolocationandothersenses 61 Whatisecholocation?............................................................................................................61 Soundgenerationandperceptioninecholocatingbats...................................................63 Echolocationcalls ..................................................................................................................64 Howdobatsavoidbeingconfusedbyotherbats’sonar? .............................................80 Auditoryadaptationsandtheneuralbasisofecholocation ..........................................80 Echolocationandbehaviouralstudiesinthewild............................................................82 Thecostofecholocationanditsconsequences..................................................................86 Othersenses ...........................................................................................................................88 Theecologyofecholocation:interactionsbetweenflight,food,andforaging habits .......................................................................................................................................90 Evolutionofecholocation......................................................................................................92 4. Torporandhibernation 97 Energybalance........................................................................................................................97 Concepts:ectothermy,endothermy,andheterothermy...................................................97 Concepts:torporandhibernation .......................................................................................99 Thecostofstayingwarm—theimportanceofbodysize ................................................99 Thephysiologyoftorpor................................................................................................... 102 vi TABLE OF CONTENTS Theecologyoftorporandhibernation ........................................................................... 104 Biologicalclocks.................................................................................................................. 109 Evolutionoftorporandhibernation .............................................................................. 111 5. Reproductionanddevelopment 113 Reproductivecycles............................................................................................................ 113 Gestation.............................................................................................................................. 118 Birth,development,andsurvival .................................................................................... 119 Infant–mothercommunicationandmaternalcare......................................................... 124 Roostselectionandmaternalforagingpatterns ............................................................ 125 Matingbehaviour ............................................................................................................... 126 6. Roostingandfeedingecology 137 Roostingecology ............................................................................................................... 137 Feedingecology .................................................................................................................. 154 Informationtransfer............................................................................................................ 163 Interactionsbetweenhabitat,food,foraging,socialstructure,andbehaviour ......... 165 7. Migration,socialstructure,andpopulationstructure 175 Migration.............................................................................................................................. 175 Navigation........................................................................................................................... 179 Socialandpopulationstructure ....................................................................................... 180 8. Biogeography,macroecology,communityecology,andtheinteractions betweenbatsandotherorganisms 195 Biogeographyandmacroecology .................................................................................... 195 Communityecology............................................................................................................ 211 Predatorsandprey ............................................................................................................ 220 9. Conservation 239 Whyconservebats?............................................................................................................ 239 Anoverviewofcurrentstatus.......................................................................................... 239 Identifyingandpredictingconservationneeds ............................................................ 241 Threatstobats..................................................................................................................... 243 Fromthenegativetothepositive:howtoconservebats ............................................. 266 Batconservationorganizations ....................................................................................... 284 References 285 Index 319 Preface Preface to the first edition advances in almost all areas of bat biology and most chapters have required a major rewrite and Thisbookisaimedprimarilyatundergraduateand expansion. In some areas developments have been graduate students wishing to learn about bats, but so significant I have created new chapters. I have also aims to show how a study of one group of alsoincludedamajorchapteronconservation. My animalscancontributetoawiderunderstandingof interestinbatsbeganinconservationandithasalso theprocessesthatshapethenaturalworld.Itthere- been responsible for the shift in my own research forehastwomainobjectives.Thefirstistogivean from biomechanics to bat ecology. Bats are under accountofthebiologyoftheworld'sbats,emphasiz- threat from habitat destruction, degradation and ingthoseaspectsthatareuniqueorhighlyadapted, fragmentation, from climate change, and from the notably flight and echolocation. The second objec- manyotherpressures humansbringtobearonthe tiveistoillustrateprocessesandconceptsofbroad planet. They therefore need our help. As a large, biological relevance, many of which are major diverse, and widespread group of animals, their themes in current research. The coverage is broad, value as indicators of environmental health has but by no means comprehensive. I have tried to long been evident to biologists and has recently make the book accessible to the growing number been more widely recognized. Although the book of bat enthusiasts in all walks of life, by giving isbroaderinitsscopethanitsparentitisnoweven some relevant background to what I imagine are moredifficulttobecomprehensive,soIapologizeif themoredifficultsectionsandbyexplainingtermi- your favourite subject gets too little attention! It is nologyandprinciplesthatmaybeunfamiliar. increasingly difficult to do justice to the technical complexity of many areas of study in a book of such breadth. I have tried to walk that fine line: Preface to the second edition givingyousufficientdetailtoproperlyinformwith- In this updated andexpanded versionmy aimhas outgettingyoumiredinthedetail.Inbringingyou notchanged,butthestateofbatbiologyhas!Scien- uptodatewiththekeyadvancesinourknowledge tificandpublicinterestinbatshasgrownconsider- and understanding of bats, I hope I have also illu- ably in recent years and the scientific literature on strated some of the major changes in the way we bats has exploded. There have been significant thinkaboutandstudythenaturalworld. This page intentionally left blank Acknowledgements MostofthedrawingsofbatsarebyTomMcOwat,a IamverygratefultoRogerButlinforintroducing few by Lucy Hammond—all are superb and make metothepowerofmoleculargeneticsandtoMan- thebookatreatfortheeye.Thecolourphotographs uelRuediforcastinghisexperteyeoverthegenetics werealltakenbyMerlinTuttleandIamgratefulto sectionsofthebook. Bat Conservation International for making these MywifeKateandourdaughterAnnedidmostof available.Manyoftheblackandwhitephotographs thehardworkofcompilingandcheckingtherefer- aremyown,andwheretheyarenot,thephotogra- ences—ataskIwasverygratefultopasson!Again, phersaregratefullyacknowledgedinthelegends. ifmistakesremaintheyaremine. Chloe Bellamy, Anita Glover, Elena Papadatou, HelenEatonatOUPhasbeenamodelofpatience, Kirsty Park, Chris Scott, and Paula Senior read efficiency, and helpfulness and it has been a plea- manyorallofthechapters—manythanksforyour suretoworkwithher.IanShermanatOUPhasalso comments,correctionsandencouragingwords.Any been very patient—Ian, I hope this long overdue remaining failings are of course entirely mine! bookisworththewait. Thankstoallofthemandtootherpastandpresent Most of all, thanks to Kate for her unwavering members of my research group for the hard work support! andfunwe’vehadwhilestudyingbats.
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