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DUNGBEETLES,MONKEYS,ANADMASEZEODNDISPERSALINTHEBRAZILIAN By KEVINAVULINEC ADISSERTATIONPRESENTEDTOTHEGRADUATESCHOOLOFTHE REUQNUIVIERRESMIETNYTSOFFFOLROTRHIEDADEINGRPEAERTOIFALDOFCUTLFOIRLOLFMEPNHTILOOFSTOHPEHY UNIVERSITYOFFLORIDA 1999 Tothememoryofmybrothers,bloodandspirit: DavidClover 1939-1997 JohnE.Fischer 1948-1998 and Zack(TheKing) 1979-1999 ACKNOWLEDGMENTS Iwishtothankthemembersofmycommittee,CarlBarfield,ColinChapman, JimLloyd,andMikeThomas,fortheirsupport,advice,andencouragement.I especiallythankmycommitteechair,JohnSivinski,forhisenthusiasm,support,and intellectuallyintriguingdiscussions. Ithankthefollowingorganizationsfortheirfinancialsupportofmyresearch: TheEntomologyandNematologyDepartmentattheUniversityofFlorida,the FulbrightCommissionforInternationalStudies,theGahanFoundationatthe UniversityofFlorida,theCharlesA.andAnneMorrowLindberghFoundation,the FloridaCenterforSystematicEntomology,theDickinsonAwardforTropical Agriculture,theWomeninAgricultureClub,andtheJ.D.TurnerFoundation. InBrazil,IamgratefultoOConselhoNacionaldeDesenvolvimento CientfficoeTecnologico(CNPq),InstitutoNationaldePesquisasdaAmazonia (INPA),andMuseuParaenseEmflioGoeldi(MPEG).Iamespeciallythankfulto ClaudioR.V.daFonsecaofINPAforhisfriendshipandimmeasurablehelp.Ialso wishtothankPedroL.B.Lisboaforhisassistance. SpecialthanksareduePhoebeWilsonforthemaps,DavidAlmquistand FreidaAnsoanuurforsortinginsects,BruceGillandDaveEdmondsforhelpwith identifications.IamverygratefultoKenPortier,GalinJones,andVictorHuforhelp withthestatistics.IthankGinaPoseyandSidMayerfortheircomputerand programminghelp. Ialsothankmysons,ColemanandCoreyKanefortheirhelpinBrazilandthe States,andtheirnever-endingconfidencethatIwouldeventuallyfinishandgetareal job.Ithankmyfather,TonyVulinec,withlove,foralifetimeofsupport.Thanksare alsoduePetteBlanchard,JulieCandela,AlfonsoDiaz,andJoannaLambertfor continualfriendshipandencouragement(wewillsurvive).Finally,withoutthehelp andsupportofmyfieldassistant,packer,monkey-spotter,andentertainment committee,myhusband,DaveMellow,thisprojectwouldhavediedalonelydeathin Rondonia. TABLEOFCONTENTS page ACKNOWLEDGMENTS iii ABSTRACT vii INTRODUCTION 1 EvolutionofDispersalCharacteristicsofSeeds 3 OverviewofSeedDispersal 5 CommunityEcologyandSeedDispersal 8 EcologyofDungBeetles 9 DungBeetlesasSeedDispersers 13 BackgroundonMonkeysasSeedDispersers 16 DungBeetlesandDeforestation 25 ForestRegenerationandSeedDispersal 27 Objectives 28 METHODS 31 StudySites 31 Surveys 34 SeedBurialExperiments 38 RESULTS 40 BeetleSurveys 40 MonkeyCensuses 41 MonkeyandBeetleComparisons 42 BeetlesandSeedBurial 42 v CommunityAnalysis 47 DISCUSSION 88 APPENDIX1.BEETLESPECIESCOLLECTED 98 APPENDIX2.MONKEYOBSERVATIONS 99 LITERATURECITED 102 BIOGRAPHICALSKETCH 116 AbstractofDissertationPresentedtotheGraduateSchool oftheUniversityofFloridainPartialFulfillmentofthe RequirementsfortheDegreeofDoctorofPhilosophy DUNGBEETLES,MONKEYS,ANADMASEZEODNDISPERSALINTHEBRAZILIAN By KevinaVulinec December1999 Chairman:Dr.JohnSivinski MajorDepartment:EntomologyandNematology Tropicalrainforesttreeshaveevolvedstrategiestomoveseedsawayfromthe parentplant.Thesestrategiesarethoughttoreducetransmissionofpredatorsand fungaldiseasesfromparenttooffspring.Oneofthemostcommonstrategiestomove seedsisthroughfruitcarriedinvertebrategutsanddefecatedlater.Seedsembeddedin dunganddepositedontheforestfloorarevulnerabletoseedpredationbyrodents unlessburiedbydungbeetles.Inthisproject,Iexplorethedynamicsofdungbeetle- monkey-seedinteractionsandhowthisprocessmaycontributetoforestregeneration, especiallyinsecondarygrowthanddisturbedhabitat.Workwasdoneatthreesiteson theAmazonRiverinBrazil:Esta<;aoFerreiraPennainParaandReservaAdolfoDucke inAmazonas,andprivatelandnearCacaulandia,Rondonia.Ideterminedthatdifferent speciesofdungbeetleshavevaryingefficienciesatseedburial,andbeetlespecieswere rankedintoseedburialguildsbasedonscoresfromaprincipalcomponentanalysis. Thepotentialforseedstobeburiedisdependentonthedungbeetleandmonkey communitiesateachsite.Communitiesamonghabitatswithinthesamesitealso differ:clear-cutareasharboraverylowbiomassofdungbeetles,andthespeciesare differentthaneitherprimaryforestorsecondarygrowth.Secondarygrowthcontainsa subsetofthesamespeciesfoundinprimaryforest,butlowerinspeciesrichnessand abundance.Correspondenceanalysisofhabitatsandseedburialguildssuggeststhat effectiveseeddispersalandburialinsecondarygrowthdependsontheamountof disturbanceofthathabitat.Somemanagementrecommendationsaremadethatmay encouragemonkeyandbeetleseeddisperserstocolonizedisturbedareasandcontribute toforestregeneration. UNIVERSITYOFFLORIDA 1999 INTRODUCTION Rainforestsaredefinedandstructuredbytheirtrees.Theseforestsarealso dynamic;gapsarecreatedbytreefalls,allowinglighttolerantspeciestogrow,andgaps arefilledinsuccessionally,becomingprimaryforestagain(Martinez-Ramos& Alvarez-Buylla1995;Brokaw1986,1985).Regenerationintropicalforestsis dependentonseeddispersal,mostlyaccomplishedbyvertebrates.Althoughdispersal canbebywind,water,orotherfactors,intheNeotropics,45-90%ofcanopytreesand almost100%ofshrubsandsubcanopytreeshavefruitadaptedforanimaldispersal (Terborgh1983;Howe&Smallwood1982;McKey1975).InNeotropicalwet forests,only10%oftreesaredispersedbywind;incontrast,21%oftreesintemperate forestshavewinddispersedseeds(Dirzo&Domfnguez1986).IntheNeotropical rainforests,monkeysareoneoftheprimarydispersersoftreeseeds,swallowingseeds wholeatthefruitingtree,anddefecatingthemintactadistanceawayfromtheparent tree(Emmons1990;Estrada&Fleming1986;Janson1983).Primatesareestimated tobebetween25-40%offrugivorebiomassintropicalforests(Chapman1995; Terborgh1983;Eisenberg&Thorington1973).Nevertheless,morethan90%of seedsdispersedbymonkeysmaybedestroyedbyrodents(Estrada&Coates-Estrada 1991).Secondarydispersalandburialbydungbeetlesmayproviderefugiaforseeds fromrodentpredationandbeanimportantforceinrainforestregeneration.Previous workhasemphasizedthepotentialroleoftheseinsectsindispersalandseedburial 1 2 (Andresen1999,1994;Feer1999;Estrada&Coates-Estrada1991;Howard&Zanoni 1989)andingermination(Shepherd&Chapman1998). Dungbeetlesplayintegralrolesintropicalecosystemsasdecomposers,nutrient recyclers,andseeddispersers.Perturbationsoftheenvironmentchangedungbeetle communitycompositionand,often,decreasediversity(Vulinec,inpress;Halffteret al.1992;Klein1989;Moron1987;Howden&Nealis1975).Ascattleranching, logging,andconcomitantdeforestationincreaseinthetropics(anannualmean deforestationrateof0.8%between1980and1990;WorldResourcesInstitute1994), manyspeciesofdungbeetlesthatareadaptedtorainforestconditionsmayvanish. Notonlywouldextinctionofthisbeetlefaunaaffectbiodiversity,buttheimpactofits lossonsoilstructure,nutrients,predatoryphoreticmitesandnematodes,andforest regenerationisunknown. Recently,muchattentionhasfocusedontropicalforestregenerationand reclamation(Lauranceetal.1997;Wunderle1997;Laurance1991),particularlyin placesthathavebeendeforestedforcenturies,suchasareasintheNortheastofBrazil (Uhletal.1997).Insomecases,abandonedpasturesandplantationsmayregenerate backtotheoriginalforest,ifenoughintactrainforestisleftinthesurroundingarea,and ifuseofthedegradedareawaslightormoderate,e.g.notbulldozed(Buschbacheret al.1992). However,notonlymusttherebeintactforestleft,butthefaunainvolvedin ecosystemfunctionsalsomustbepreserved,includingthepollinatorsanddispersers (Pannell1989). Myresearchhastwosetsofobjectives;(1)delineatethecharactersthatmake particularspeciesofdungbeetlesgoodseeddispersalagents(dispersaldefinedaseither horizontalmovementorburial),establishanindexofseeddispersalabilities,andrank eachspeciesintermsofrelativequalityasaseeddisperser,and(2)examinewhether