THREATSTO NATIVEAQUATICINSECTBIODIVERSITYINHAWAI'IAND THEPACIFIC, ANDCHALLENGES INTHEIRCONSERVATION ADISSERTATIONSUBMITTEDTOTHEGRADUATEDIVISIONOFTHE UNIVERSITYOFHAWAI'IINPARTIALFULFILLMENTOFTHE REQUIREMENTS FORTHEDEGREEOF DOCTOROFPHILOSOPHY IN ENTOMOLOGY AUGUST 2005 By RonaldA. Englund DissertationCommittee: MarkWright, Chairperson DanRubinoff NealEvenhuis DanPolhemus AndrewTaylor TABLEOFCONTENTS ACKNOWLEDGEMENTS , ii ABSTRACT iii LISTOFTABLES vi LIST OFFIGURES viii CHAPTER 1.THEIMPACTS OFINTRODUCEDPOECILIIDFISHANDODONATAONTHE ENDEMIC MEGALAGRION(ODONATA)DAMSELFLIES OF0'AHUISLAND,HAWAI'I 1 CHAPTER2: EVALUATINGTHEEFFECTS OFINTRODUCEDRAINBOWTROUT(Oncorhynchus mykiss) ONNATIVESTREAMINSECTS ONKAUA'IISLAND,HAWAI'I 40 CHAPTER3. LONG-TERMMONITORINGOFONEOFTHEMOSTRESTRICTED INSECT POPULATIONSINTHEUNITEDSTATES, Megalagrion xanthomelasSelys-Longchamps, 1876,AT TRIPLERARMYMEDICALCENTER, O'AHU,HAWAI'I... 76 CHAPTER4.THELOSS OFNATIVEBIODIVERSITYANDCONTINUINGNONINDIGENOUS SPECIESINTRODUCTIONSINFRESHWATER,ESTUARINE, AND WETLANDCOMMUNITIES OFPEARLHARBOR,O'AHU,HAWAIIANISLANDS 91 CHAPTER5. FLOWRESTORATIONAND PERSISTENCEOFINTRODUCED SPECIES IN WAlKELESTREAM,0'AHU 125 CHAPTER6: INVASIVESPECIESTHREATS TONATIVEAQUATICINSECTAND ARTHROPOD BIODIVERSITYINHAWAI'I,THEPACIFICANDOTHERRELEVANTAREAS WITHDISCUSSION OFCONSERVATIONMEASURES 143 ACKNOWLEDGEMENTS Iwouldliketo thankthemanypeoplethathavemadethis dissertationpossible. Iespeciallywould liketo extendmythanksandwarmestgratitudetomyadvisorMarkWright, whosesenseofhumorandkeen intellectmadethis process as enjoyableas itcan be. Mycommittee,consistingofDanRubinoff, Neal Evenhuis,DanPolhemusandAndrewTaylorprovidedvaluableinsightsandadvicethroughout. Isincerely acknowledgeandappreciatethe effortsofmyentirecommitteethroughoutmytimeattheUniversityof Hawai'iatManoa.Iamalsodeeply appreciativeofNealEvenhuisandAllenAllisonfortheir encouragement,andallowingmetheflexibility topursueaDoctoratewhile beingemployedatthe Bishop Museum.Ihavegreatlyenjoyedthescientificandcultural 'ohanaattheBishopMuseumthatalways providedanidealresearchandworkingatmosphere. RobCowieandFrankHowarthoftheBishopMuseum alsoprovidedvaluablereviewsandadviceformanyofthesechapters. BishopMuseumlibrariansPatti BelcherandB.J.Shortwerealwayshelpfulintrackingdownthemanyobscurereferences.Severalkey organizationsprovided thesupportthatallowedmyresearchtotakeplace,andIthankthefollowing organizationsthatfundedthisresearch: BishopMuseum, HawaiiDivisionofAquaticResources,Nature a Conservancy,SmithsonianInstitution, andtheDelegation laRecherchePolynesiefran<;aise. Peopletoonumeroustomentionassistedmeinvariousaspectsofthefieldworkrequiredforthis wide rangingdissertation, andIgreatlyappreciatehelpfromDavidPreston,BetsyGagne, DanPolhemus,Jean Yves Meyer, BenoITFontaine, OlivierGargominy,TinaLau,StephanieLoo, BrianNaeole, Alison Sherwood,andSteveJordan. SpecialthanksgoestotheHawaiiDivisionofAquaticResources crew includingBobNishimoto,GlennHigashi,DarrellKuamo'o, JohnKahiapo, SkippyHau, BillPuleloa, and MikeYamamoto. BillDevickandBobNishimotowerealsoinstrumentalinencouragingandfundingmuch ofthisresearchaswell. LaVonneFurtadoprovidedconsistentmoralsupportduringthecrucialfinal stages ofthisjourney.Iamalsoprivilegedtohaveincrediblysupportiveparents andawonderfulfamily, without whomIwouldhaveneverattemptedthis work. Iwouldliketodedicatethis dissertationtomywonderful parents,StanleyandMarjorieEnglund. 11 ABSTRACT AlthoughthedeclineinnumbersanddiversityandthreattonativeinsectsintheHawaiianIslandsis widely recognizedbyfield scientiststherehasbeenlittleprogressineitherdocumentingtherealdeclineofnative species,orindemonstratingspecificcausesoftheoveralldeclineofthesespecies. Additionally,few conservationactions toeitherrestorepopulationsormitigateactualthreatstonativearthropodshavebeen mentionedintheliterature.Thefollowing chaptersexamineseveralassessmentsofrelevantaquaticsystems andthenativeaquaticinsectsdwellingwithin,wheretherehaseitherbeenaperceivedorrealdeclineofthese nativeHawaiianaquaticarthropodsbecauseofthreatsfrom invasiveorintroducedspecies. Thelargeadaptiveradiationoftheendemicnativedamselflies(Coenagrionidae: Megalagrion) inHawai'ihas receivedconsiderableattentionandstudysinceatleastthe 1880s.EndemicMegalagrionareinmanyways reflectiveofagreatlossbecausetheyarelargelynowfound inremoteupperheadwaterareasofstreams,yet theyalsorepresentthehopeofpreservinghighlydiversefreshwaterecosystemsfoundthroughoutthe Hawaiianarchipelago.Thefirsttwochaptersofthisdissertationexaminetheimpactsoftwodifferingtaxa ofintroducedfishonHawaiianMegalagrion, Poeciliidae(livebearersormosquitofishfamily) and Salmonidae(trout).Theeffectsofeachfish speciesonnativeaquaticinsectsdependedmainlyonthe invasivestatusofeachgroup; forexample,Chapter1(Englund 1999)examinestheimpactsofintroduced poeciliidsonnativedamselflies. Damselflieswerecompletelyeliminatedontheislandof0'ahuwherever speciesinthehighly invasivemosquitofishfamily werefound, andonlyremnantpopulationswerefound in highelevationslackingintroducedfish. Chapter2(EnglundandPolhemus2001)examinestheimpacts of thenon-invasiverainbowtrout(OncorhynchusclarkI) onMegalagriondamselflies. Damselfliesandallother nativeaquaticinsectswerenotfoundtobeharmedbytroutintheuppermostelevationsofKaua'istreams wheretroutreproducenaturally, andevenhadmorerobustpopulationsthaninsomenearbynon-trout containingstreams.Thelackofimpactsonnativedamselfliesbyalarge,generalistpredatorsuchas rainbowtroutpointedoutaseemingparadox.Whereas thesmallbutubiquitousmosquitofishappearsto havecompletelydevastatednativeaquaticfaunawhereverithasbeenintroducedoutsideofitsnaturalrange, iii trout,becauseoftheirrestrictedrange andsmallerpopulationsizeshavehadminimal,ifanyimpactson nativeinvertebratesinHawai'i. Becauseintroducedfish specieshavecausedeithertheextinctionorsevererangecontractionsofMegalagrion damselfliesinHawai'i,long-termmonitoringoftheremnantpopulationshasbecomenecessarytopreserve theseremainingpopulations. Chapter3(Englund2001)providesacasestudyinboththemonitoringand preservationofaremnantO'ahudamselflypopulationnowfound inonly95moffishless streamatthe TripIerArmyMedicalCenter. Chapter3alsoprovidesseveralharrowingexamplesofhowthisspecieswas nearlybeeneliminatedinthepast10years throughaccidentsandmismanagement. Notonlyaretheendemic Megalagrionnowmissingfrom alllowlandareasofO'ahu(withtheexceptionoftheTripIerpopulation), lowlandaquaticinsectdiversitythroughoutO'ahuisataremnantstatus,andbiodiversitysurveysfornative aquaticinsectsinthePearlHarborwatershedsinChapter4(Englund2002)indicatedanearabsenceof nativeaquaticinsectsinthesefreshwaterhabitats.LowerPearlHarborwatershedsweredocumentedtohave lostmanynativeaquaticinsecttaxasuchas allnativeHeteroptera,damselflies,Coleoptera,andmany Dipteraspecies,whileintroducedinsectspecieswereabundant. Avarietyofconservationmeasureshavebeensuggestedtoeitherrestoreormaintainthecurrentlevelsof freshwaterbiodiversityinHawai'i.InChapter5(EnglundandFilbert1999),thecaseofsignificantly increasingandrestoringstreamflow inaformerlydivertedstreamwasexaminedtodeterminewhetherthis factoralonewouldleadtoarestorationofnativeaquaticspecies.Itwasfoundthatmerelyincreasingstream flow byitselfwasnotenoughtoridthestreamofanyalienaquaticspecies,infact, severalnew nonindigenousaquaticspeciesbecameestablishedafterstreamflows wereincreased.TheresultsofChapter 5confirmthatanintegrated,balancedandpossiblydrasticapproachwillberequiredtomaintainandpreserve Hawai'i'snativeaquaticinsectfauna. Awide-varietyofconservationmeasuresintheHawaiianarchipelago willbeneededtomaintaincurrentbiodiversitylevels,andalsohopefullyrestorenativefreshwater biodiversityinselectedareas. iv ToputtheHawai'iproblemintoperspective, abriefreviewoftheimpactsofinvasivespeciesonnative insectsinothertropicalareas isprovidedinChapter6.Thisreviewchapteralsoprovides asynthesisofthe problemfacingHawaiianfreshwaterinsectsandotherterrestrialarthropods inHawai'iandelsewheredueto invasivespecies,andhowtheHawaiiancasestudyofinvasivespeciesimpactshasmanyparallelsto other vulnerablebiotas.Finally,drawingonamixedrecordofpastmistakesandsuccessesinHawai'iand elsewhere,somepotentialpracticalconservationmeasuresintendedtopreserveandrestoreendemicisland aquaticinsectsareprovidedinChapter6. v LISTOFTABLES Table1.1.ExtinctionsofMegalagrionspeciesinsurveyed0'ahuaquatichabitatssince 1936 8 Table 1.2.RemnantnativeMegalagrion speciesfound inO'ahustreams,tributariesinparentheses,and relativeabundanceinstreamareascontainingnativedamselflies, (rare(R)=<3individualscollectedor observed); (moderatelycommon(C)=<:3individualscollectedorobserved) 9 Table 1.3.BiogeographicstatusandHawaiianIslanddistributionofaquaticMegalagrion andintroduced OdonataspeciesfoundonO'ahu 11 Table2.1. AquaticinsectspeciesandnativeorintroducedstatuscollectedateachKaua'istream 50 Table2.2.Totalnumberofaquaticspeciescollectedduringbenthic,drift, andaerial(general)collections duringthisstudy 51 Table2.3PresenceorabsenceinsurveyedK6ke'eStateParkstreamsofnaturallyreproducingandstOcked rainbowtroutandnativeMegalagriondamselflies 57 Table2.4.Geographicoriginandterrestrialoraquaticstatusofpreyitemsfound in80K6ke'etrout stomachs, 1997-1999 60 Table2.5. Summarynumbersandpercentfrequency ofnativepreyitemsofspecialconcerncollectedin rainbowtroutstomachsduringthis study,comparedtothenumberoftaxacollectedperstream 61 Table4.1. Summaryofthenativeornonindigenousstatusandtotalnumber(percent)ofaquaticspecies found inPearlHarborestuarinehabitats 98 Table4.2. Geographicsource(yearofintroduction)andknown (orprobablyknown)modeofintroductionof nonindigenousspeciesofaquaticmacrofaunafoundinPearlHarborstreamsandestuaries 107 Table5.1.Therangeandmeanwatervelocities(±standarderror)recordedintransectsdownstreamof WaikeleSprings 131 Table5.2.Introducedandnativespeciesfound inWaikeleStream,Oahuin 1993and 1997-1998from 250 maboveWaikeleSpringsdownstreamtoconcreteweir. OahuintroductiondatesfromBeardsley (1980), Devick(1991a), Cowie(1995), Polhemus & Asquith(1996), Randall(1996), Cowie (1998). ....................................................................................................................................132 vi Table6.1. ExtinctionstatusofnativeinsecttaxaintheHawaiianIslands thathaverecentlyhadtheir conservationstatusexamined,to lowesttaxonomicresolution 149 Table6.2.SuccessfullyeradicatedinvasiveanimalspeciesintheHawaiianIslands 161 vii LIST OFFIGURES Figure 1.1.Limnologicaldivisions onO'ahudefinedforthepurposesofthis study 5 Figure 1.2.Statusofstreamandwetlanddwellingdamselfliesontheislandof0'ahu. E=Extincton0'ahu .......................................................................................................................................8 Figure 1.3.ElevationaldistributionforHawaiianMegalagriondamselfliesandintroducedpoeciliidfish for streamsenteringKane'oheBay,O'ahu 12 Figure 1.4.ElevationaldistributionforHawaiianMegalagriondamselfliesandintroducedpoeciliidfishfor streamsenteringnorthernwindwardO'ahu 13 Figure 1.5.ElevationaldistributionfortheHawaiianMegalagriondamselfliesandintroducedpoeciliidfish forstreamsenteringPearlHarborandleewardO'ahu 14 Figure 1.6.ElevationaldistributionforintroducedIschnuraramburiiandIschnurapositadamselfliesand introducedpoeciliidfishforselectedKane'oheBayandPearlHarborstreams,O'ahu 18 Figure2.1. StudyareaofsampledK6ke'eStateParkStreams,Kaua'iIsland,Hawai'i 45 Figure2.2SummaryofaquaticspeciescollectedinK6ke'eStateParkstreamsfromallsamplingmethods combined(generalsampling,Malaisetraps, drift,benthicsamples) 52 Figure2.3. Summaryofallinsectspeciescollectedfromrainbowtroutstomachs(n=80) andtheir terrestrialoraquatic,andnativeorintroducedstatusinK6ke'eStateParkStreams 52 Figure 2.4. Meandensity by streamforthetwomostimportantconstituentsofbenthic (Surber) samples, thecaddisfly C. pettitiandmidge C. bicinctus 53 Figure2.5. SummarygraphofnumberofaquaticspeciesinKaua'iStreamsandthepresenceorabsenceof naturallyreproducingtroutineachstream;Lumaha'iandHanaleihaveneverbeenstockedwithtrout. ..................................................................................................................................... 53 Figure2.6. ThesixnumericallymostabundantaquaticinsecttaxacapturedindriftsamplestakeninK6ke'e stateparkstreams 58 Figure3.1. MapofO'ahu,Hawai'ishowinglocationsofcurrentandhistoricrecordsfor Megalagrion xanthomelas(fromEvenhuis eta1., 1995) 78 viii Figure 3.2. TAMCMitigationpondspriorto drainage, February 2000 79 Figure3.3. Megalagrion xanthomelascapturesatTAMC stream,May 1997-June2000 82 Figure 3.4. Megalagrion xanthomelascapturesatTAMCmitigationpondsfrom May 1997-February2000. ..................................................................................................................................... 83 Figure 3.5. Oviposition scars on waterlilies atthe TAMC mitigationpondsJuly 1997-February2000.. 84 Figure4.1. MapofPearlHarborwithsamplinglocations 95 Figure4.2. Numberofspeciesbystreamandnativeornonindigenousstatusforcombinedaquaticfauna found inestuarineregionsofPearlHarbor 98 Figure4.3. Nativeornonindigenousstatusoffish speciesandtotalnumbersfound atdifferentsalinity levelsinPearlHarborestuaries 101 Figure4.4. Nativeornonindigenousstatusofaquaticinsectsatvaryingelevations onO'ahu: upperHalawa datafromPolhemus(1994),upperWaikeledatafromEnglund(1993) 112 Figure5.1. WaikeleStreamstudyarea 128 ix
Description: