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BIOLOGYANDCONSERVATIONOFTHEHARPYEAGLE INVENEZUELAANDPANAMA BY EDUARDOALVAREZ-CORDERO ADISSERTATIONPRESENTEDTOTHEGRADUATESCHOOL OFTHEUNIVERSITYOFFLORIDAINPARTIALFULFILLMENT OFTHEREQUIREMENTSFORTHEDEGREEOF DOCTOROFPHILOSOPHY UNIVERSITYOFFLORIDA 1996 ACKNOWLEDGMENTS Manyprivateandgovernmentinstitutionsandagreatnumberofindividuals contributedtotherealizationofthisstudy. However,beforeanyandall,Iwishto recognizeandthankLucieE.Zaugg,friendandwife,forthecommitmentandloving supportanddedicationinvestedtoensurethatourfamilysurvivedtheselongyearsas webecamestudentsagain.Webothknewthatitwouldbeaperiodfullofgrowth, sacrifices,andtrials.LuciehasdonemorethanhershareeverytimethatIdisappeared intothejungle(oncefor3months),andhastolerated(barely)havingmyworkofficeat home.Ourtwosons,JorgeDavid(12)andJuanMichael(9),havegrownwiththe projectandarewaitingandwonderingifthereisalifeafterthePh.D. WecametoGainesville,Florida,thankstotheencouragementofPoly,Dr. EleanorStorrs,whoconvincedDr.KentRedfordtogivemeanaudience.HeandDr. JohnRobinsonprovidedmewiththeopportunitytobecomea"Gator,"aftera10year sabbaticalfrombeingan"Aggie,"andIthankthemforit.Atvarioustimesduringthe difficultadjustmenttouniversitylife,wordprocessors,andeconomicchallenges,their ProgramforStudiesinTropicalConservation(PSTC)cametotherescue(evenwhen theyknewthatIwasnotfromMexicoorBrazil). Dr.MichaelW.Collopyperhapsdidnotdeservetogetstuckwithmeandthis project,butsuchislife.Heapproachedthischallengewithhisalwayspositiveand generousattitude.HerewasaLatino,withZEROexperienceworkingwithraptors, sayingthathewouldtacklethebiggestoneofthemall.Hehasbeenapatientmentor, answeringmyeveryquestion,carefullyeditingmydocuments,andkindlyrespectingmy frequentignoranceabouthisfieldofexpertise.Hewasgenuinelycommittedtoseeme ii through,evenafterrealizingthatfundingwouldbeverydifficulttoobtain.Icannot thankhimenoughfortrustingmetogetthejobdone,andfortheconstant encouragementduringthemanymonthsittookmetoshapethisdissertation.Theone thingIknowthatImustdotoshowhimmyrespectistoturnthisdocumentintopeer- reviewedpublications. Drs.MelvinE.Sunquist,SusanK.Jacobson,MarianneC.Schmink,andRichard A.KiltieservedonmycommitteeandwereextremelysupportivewhileIpreparedthis dissertation.Theyeachprovidedadifferentinsightandcarefuleditingtoimprovethis document.Iappreciatetheirpatienceandinterestinthisprojectthatpotentiallyhad many"looseends"andfewguaranteesofbeingattainable.Ienjoyedtheopportunityto takeclassesfromDr.SchminkandDr.Jacobson;Ilearnedmuchfromthose experiencesandbothofthesesuper-teachers. IrecognizethesupportoftheUniversityofFlorida,anditsDepartmentof WildlifeEcologyandConservationfortheopportunitytotrainandlearnonthiscampus, fortheequipmentloans,thewonderfullibraries,andflawlessaccesstotheInternetso thatIcouldtracktheHarpyEaglesfrommyPC. In1988,1visitedThePeregrineFund(Boise,Idaho)andmetDr.BillBurnham, Presidentofthatorganization.WediscussedmyplanstoworkwiththeHarpyEagle. AfterImovedtoFlorida,hestayedintouchandoftenofferedhissupport.In1991,as soonasIwasreadytogointothefield,itbecameevidentpromisedfundingwasnot forthcoming posinganadditionalchallengeforthisproject.Througha"gentlemen" , agreementbetweenDr.CollopyandDr.Burnham,Iwaspresentedwiththeunique opportunityofdevelopingthefieldprojectfortheHarpyEagleConservationProgramof ThePeregrineFund.Theyemployedmeandsupportedalltheneedsoftheproject. Theyhavetreatedmeasaprofessionalandafriend, whilealsoallowingmethe latitudeofbeingastudent.Icouldnothavehopedforanythingbetter,butthe iii Burnhamsalsohaveprovidedmyfamilywiththeconstantassurancethatevenifwe werenotinBoise,webelonged.Thewholeheartedandongoingcommitmentofthe "PeregrineFundFamily"tosupportthisprojectislargelycreditedforwhatwehave beenabletoaccomplishsofar.Iextendmygratitudetoallmycolleagues,coworkers, directors,andthemanymembersthatcontinuetosupportthiswork. In1989,IbegantrainingayoungVenezuelanwhoclimbedthefirstactiveHarpy EaglenestthatDr.CollopyandIfoundduringthepreliminaryfieldseason.Eladio RafaelAlvarezwassponsoredin1992byThePeregrineFundasafull-timeField Assistanttothisstudy.Hisdedicationandcontributiontoallaspectsofthefieldwork haveinagreatmeasuremadethisstudypossible.Rafaelistheworldexpertonthis species,havingseenandhandledmoreeagles,andhavingfoundandclimbedmoreof theirneststhatanyoneelse;wheneverIwrite"I",pleaseread"RafaelandI". IwishtomakespecialmentionoftheCenterfortheStudyforTropicalBirds, Inc.,anditsDirector,JackClinton-Eitniear,whofundedmyfirstfieldtripin1989.Ithank themforbelievingitcouldbedone! InPanamaweworkedundera5-yearcooperationagreementestablishedin 1991betweenoursponsoringorganization,ThePeregrineFund,andtheProtected AreasbranchoftheNaturalResourcesInstitute(InstitutodelosRecursosNaturales Renovables-INRENARE).Wemadenumerousfriendsinthiscountry,wherewealways feltathome.IndraS.Candanedowasaconstantcollaboratorandservedasthe nationalcounterparttoourinvestigation.KarlaAparicioandBelkysJimenezalsogave ussupport;Karlacollectedvaluableinformationduringherinternshipwiththeproject. JacoboLacsprovidedmanyimportantinsightsandlogistics,andleadustoanestin theAtlanticBasin.WealsothankourcollaboratorsinANCONfortheircooperation.Dr. NathanGaleandhiswifealsodedicatedmanyhourstohelpusanddemonstratedtheir commitmenttohelptheeagles.Wededicateaspecialmentionofgratitudetothe iv peopleoftheDarien,particularlytheparkrangersheadedby"SrChicho"whoadopted usandtookuseverywhereinatimelyandsafefashion. InVenezuelaourresearchprogramwasbackedbyCVG-EDELCAandthe VenezuelanAudubonSociety(SCAV-SociedadConservacionistaAudubonde Venezuela),andauthorizedthroughannuallyrenewedResearchPermits(Lisenciade CacehaCientifica)issuedbyPROFAUNA,theWildlifeProgramsOfficeoftheMinistry fortheEnvironment(MARNR-PROFAUNA.MinisteriodelAmbienteydelosRecursos NaturalesRenovables,ServicioAutonomoProgramasdeFauna).Since1988, EDELCA'sstaffhavesupportedthisinvestigationofHarpyEagles.Thiscompanyhas providedahousingfacilitytobasetheproject,spaceforalargeflight-pentohold captiveeagles,technicalandlogisticassistance,andeventheoccasionaluseof helicopters. ThemanagementoftheVenezuelangovernment'sregionaldevelopment corporation(L.Camposinlitt.,November30,1995),CorporationVenezolanade Guayana(CVG),grantedaccesstoanextensivedatabaseonregionalnatural resourcesheldbyTECMIN(CVG-TecnicaMinera).Wethankthemforthisprivilege. WeacknowledgethelogisticsupportreceivedfromCVG-FerromineradelOrinoco, particularlyfromtheiraviationdepartment. OurfriendsinGurideserveaspecialmention,fortheirconstantencouragement andtimelyhelp--inparticularmycoworkersatEcologia.Teo,Martin,Asterio,Leon,AN, whostilltreatmeasfamily.TheBalbasfamilywasourhotelduringthefirstyear, providing5startreatmentforallthenumerousgueststhatgotinvolvedwiththisproject. LuisA.Balbasasbeenaconstantcompanioninthiswork,basicallysolvingany problems.Iregretthathehasnotbeenabletoseemuchofthewildeagles,buthehas doneanexcellentjobcaringforthelivebirdswehaverescuedandbroughttoGuri.I alsoowehimforallthetimehehasspentadministratingtheprojectandproviding v guidancetoRafael. OrlandoBorromeintroducedmetotheuseofGPSandprovided criticalsupporttoourfieldsurveyinVenezuela. Fromthestartofthisstudy,cooperationwithagroupofForestryManagement UnitswascrucialtotheinvestigationofHarpyEagles.AlongtheImatacaRangewe workedwithSOMAGUA,CODEFORSA,MaderasNuriaI,INTECMACA,Aserradero HermanosHernandez,andMADERORCA.IntheSupamoRiverBasinourresearch wassupportedbyAserraderoMatamoros,AserraderoElManteco,andAserradero Yocoima.LuisMatamoros,andhiswifeVerenice(mysister),wereamongthefirstto enlistonourbadwagonandhaveprovidedallsortsofsupport. TheFuerzasArmadasdeCooperation(FAC),theNationalGuardinVenezuela deservespecialmentionfortrackingdownthehuntersthatshottwooftheeaglesthat weweremonitoring,andmoreimportantly,forretrievingNASA'ssatellitetrasmittersso thatwecouldusethemagain.Misioncumplida! Numerousprivateranchesallowedustoconductresearchandprovidedlogistic supportandimportantbackgroundinformationonthenestswithintheirproperties. We aregratefulforthecooperationandsupportreceivedfromalloftheseentitiesandtheir representatives. TocompileandupdatetheexistingrecordsfortheHarpyEagleweworkedin MexicowithEduardoInigo-EliasandGuidoRhar;inCostaRicawithSusanWilsonand BelkysJimenez;inColombiawithCesarMarquez;andinEcuadorwithPeterJennyand MauricioGuerrero.WebeganworkinPeruwithRenzoPiana.FernandoAntonini, anotherstudentfromthatcountry,embarkedunderourguidanceonapreliminaryfield surveyoftheTambopataregionin1996.InGuayana,wecollaboratedcloselywithNeil Rettigwhohadalreadyaccomplishedanimpressivefieldsurveyandproducedfilmson theHarpyEaglesinthatcountry.SaraGilbertagreedtocollectrecordsforSuriname duringherstayinthatcountry.SteveMcGehee,arespectedfriendandcolleaguehas vi providedanendlessstreamofpricelessreferencesthatonlyhecanlocate.Mike McMillianvisitedthefieldprojectinVenezuelatodeliverneededequipmentandprovide additionaltelemetrytrainingtoRafael. WeweretrainedtoworkwiththeeaglesbyDr.DavidEllis(NBS,Patuxent,MA), oneofthepioneeringexpertsinthefieldofsatellitetelemetryappliedtolargeavian species.Heprovidedvaluableadviseandencouragement,servingastheinvisible committeememberthateverygradstudentshouldhave.Between1992-96,Dr.Ellis arrangedthesupplyofARGOS-TIROSsatellitesystemtransmitterstoourproject throughaworldwideeffortfundedbyNASA;fouradditionaltransmitterswere purchasedbyThePeregrineFund(1993-95).Dr.PaulHowie,ofMicrowaveTelemetry, notonlyprovidedthebestsatellitetransmittersavailable,butalsogaveusthebest personalizedattentionandadvise.Irecognizethelargecontributionof NASAin supplying10satellitetransmitterstodate,andtheconstantandreliablesupportofthe peoplethathandletheincomingdata,particularlyCharlieandJon. Weacknowledgethesupportofthefollowingcompanies:TrimbleNavigation loanedtopofthelineGPSsurveyingequipmentforthecollectionofdatalayersusedin ourGeographicalInformationSystem.OursGISwasdevelopusingARCVIEW(verion 2.1byESRI)donatedtoThePeregrineFundbyEnvironmentalSystemsResearch Institute(Redlands,CA). HortonManufacturingCompany,Inc.,(Tallmadge,Ohio) donatedacrossbowusedtogetourclimbingropesoverthenesttrees;thisavoided thedamagedonetotreeswhenclimbingwithspikesandallowedustousesaferrope climbingmethods.AVINETrecentlydonatedPesolascalestotheproject. Dr.StefanGorzulaandGlendaMedinadeservecreditforgivingmethefirst trainingIeverreceivedinFieldBiology.Stefanhasbeenavaluableandsupportive colleaguesinceImethimintheearly1980s.Glendaintroducedmetotheworldof birdsandfieldtechniques.Dr.LeeHarperalsocontributedtomyearlytraining. vii Dr.OmarLinaresopenedthedoorsofhislaboratoryattheUniversidadSimon Bolivartohelpprocessandidentifythepreyremainscollectedfromnests.Oneofhis studentsassistedinthistask. NeilRettig,theotherworldexpertonthespecies,hasdemonstratedaLatino likeheartandwillingnesstoshare.Heprovidedmanyimportantsuppliesandthe crossbowtoshootlinesthatenabledustoclimbthenests.Healsowaskindenoughto guidemeintoGuyanaandshowmehisnests.Thesupportofhispartner,KimHayesis alsorecognized,aswellasthefundsprovidedtotheprojectbyNeilRettigProductions. WeoweourlivestoLuisEnriqueArnal,Kike,masterfulclimberwhotrainedus toreachthenests.ALWAYSonrope!SAFETYfirst!IrememberwhenIwasfirstgiven enoughclimbingropeandnotraining,andtoldtojustdoit!Also,Kikehasbeenthe project'sphotographersinceImethimonaCuipointheDarienin1993. Imustmentionagroupoflong-timefriendswhoadoptedthecauseofthe eaglesandpouredtheirheartsintomakingthisprojectareality.PeterE.Kung,for gettingmehookedonfieldbiology>20yrago,teachingmetoworkwithbirds,andfor histotaldedicationtotheproject'ssuccess.HealonedeservescreditfortheGPS surveys,anenormousaccomplishment,andforbelievingthatwecouldcreateaGISon ashoestringbudget.ZanMerrillprovidedusefuladviseandunendingsupport throughoutthestudy,alwayssharingthevision.GustavoA.MartinezG.dedicatedhis alltotheneedsoftheproject,servingasthein-countryclearinghouseforthe PeregrineFundduringmanyofourtrips.GusandElba,hiswife,pouredtheirkindness andhumoronmyriadsofpeopletomakesurethattheysupportedourgoals.Guswas instrumentalinsecuringtheGISinformationandbasemapsforthestudy.Hearranged planeandhelicopterrides,providedofficesupport,andevengaveourpresentationat ascientificmeetingwhenwecouldnotbethere.Milardosdegracias!!Gusalso viii traveledmanytimestoFlorida,spending nightsanddaysmakingmyPCrunflawlessly sothatIcouldcompletethisportionofthestudy. MyfamiliesinVenezuelaandCanadawerealwaysbehindmeduringthis endeavor. InCaracas,myparentsMargotandFelipe,andmysisterBelen,havebeen aconstanthelpandhaverunmanyerrandstohelpmeaccomplishthiswork.My motherbasicallydonatedherusedcarfortheuseoftheproject.Her1962FordFalcon madeinnumerabletripsintotheloggingtrailsintherainforestuntilitwastotaledinan unfortunateaccident.MybrotherCesarAlvarezhasbeenextemelysupportive, and alsoarrangedimportantlogisticcooperationfromCVGaffiliatedcompaniesinthe GuayanaRegion.RuthandFerriesinAlbertaalsosupportedthisaccomplishmentin everywaypossible. Finally,IacknowledgethewonderfuleffortsofM.J.Schaerinformattingand pullingtogetherthedraftsandfinalversionofthisdissertation,alwayswithapositive, friendly,andprofessionalattitude. ix TABLEOFCONTENTS page ACKNOWLEDGMENTS » LISTOFTABLES xiii LISTOFFIGURES xiv ABSTRACT xvi CHAPTERS 1 1 GENERALINTRODUCTION 1 Introduction 1 Background 2 WhyStudytheHarpyEagle? 3 RelatedResearch 6 FactVersusFolkloricNotions 8 InstitutionalSetting 9 OrganizationoftheDissertation 11 LimitationsofthisStudy 13 2 GEOGRAPHICANDECOLOGICALRANGEOFTHEHARPYEAGLE 14 Introduction 14 TheImportanceofHistoricalandRecentRecords 16 TheLandscapeApproachtoHabitatEvaluation 17 Methods 21 ProjectAreasandSiteSelection 21 SourcesandSearchforRecords 24 FieldSurveysinVenezuelaandPanama 27 HabitatEvaluation 30 ExplorationsinOtherCountries 31 Results 31 HistoricalandRecentRecordsinVenezuela 31 NestSurveyinVenezuela 36 HistoricalRecordsinPanama 39 NestSurveyinPanama 41 RecordsfromOtherCountries 42 Discussion 47 x