disturbance to coral reefs in aceh, northern sumatra

DISTURBANCE TO CORAL REEFS IN ACEH, NORTHERN SUMATRA: IMPACTS OF THE SUMATRA-ANDAMAN TSUNAMI AND PRE-TSUNAMI

DEGRADATION

BY

STUARTJ.CAMPBELL,1MORGANS.PRATCHETT,2AJIW.ANGGORO,3RIZYAL.ARDIWIJAYA,1NURFADLI,4YUDIHERDIANA,1TASRIFKARTAWIJAYA,1DODENTMAHYIDDIN5,AHMADMUKMININ,1SHINTAT.PARDEDE,1EDI

RUDI,4ACHISM.SIREGAR3,andANDREWH.BAIRD2*

ABSTRACT

TheSumatra-Andamantsunamiof26December2004wasthefirsttooccurinareasforwhichgoodecologicaldataexistedpriortotheeventandconsequentlyprovidedauniqueopportunitytoassesstheeffectsofthistypeofnaturaldisturbanceintropicalmarineecosystems.Lessthan100daysaftertheeventwevisited49sitesoncoralreefsinnorthernAceh,Indonesia,allwithin300kmoftheepicentre,todeterminethenatureandextentoftsunamidamageandpre-tsunamidisturbance.Reeffishdiversityandabundancewerealsoassessedinrelationtotsunamiimpactandexistingmarineresourcemanagementregulations.Atthesesites,theinitialdamagetocorals,whileoccasionallyspectacular,wassurprisinglylimitedandtrivialwhencomparedtopre-existingdamagemostprobablycausedbydestructivefishingpractices.Theabundanceofup-turnedcoralswashighlydependentonhabitatandlargelyrestrictedtocoralsgrowinginunconsolidatedsubstrataatdepth,afeaturewebelieveuniquetotsunamidisturbance.Otherevidenceoftsunamidamage,includingtheabundanceofbrokencoralsandrecentlykilledcoralswaspatchyandvariedunpredictablybetweensites:reefaspect,geographiclocationandmanagementregimehadnosignificanteffectonthesevariableswiththeexceptionofbrokenlivecoralswhichweremoreabundantatlocationswherethetsunamiwaslarger.Interestingly,therewaslittlecorrelationbetweendamagevariables,suggestingthetypeofdamageobservedwasstronglyinfluencedbywhichcoralswerepresentataparticularsiteordepth.Incontrast,reefconditionwasclearlycorrelatedwiththemanagementregime.Coralcoverwasonaverage2-3timeshigheronreefsmanagedunderthetraditionalAcehnesesystem,PanglimaLaut,andinthePulauRubiahMarineParkwhencomparedtoopenaccessareas.Turfalgaeandcoralrubblewere2-3times

1TheWildlifeConservationSociety,MarinePrograms,Bronx,NewYork10460,USA2ARCCentreofExcellenceforCoralReefStudies,JamesCookUniversity,Townsville,Queensland4811,Australia3InstitutePertanianBogor,Bogor16151,Indonesia4UniversitasSyiahKuala,BandaAceh,NAD,Indonesia5RubiahTirtaDivers,Ibioh,Sabang,NAD,Indonesia*Correspondingauthor.Tel:+617-47814857,Fax+617-47251570,E-mail:[email protected]

No. 544. Ed. Stoddart, D.R. July 2007. Tsunamis and coral reefs

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moreabundantinopenaccesssitescomparedwithmanagedareas.Theseresultsareconsistentwithahistoryofdestructivefishingpractices,suchasbombingandcyanidefishinginopenaccessareas.Coralreeffishabundanceanddiversitydidnotdifferamongmanagementzones,despitethefactthatPulauRubiahMarineParkhasbeenclosedtofishingfor10years.However,therewereconsistentdifferencesinthestructureofthereeffishassemblagesamongthesezones.Forexample,thenearabsenceofchaetodontidsatopenaccesssitesisprobablytheresultoflowcoralcover.ThehighabundanceofscaridsandacanthuridsintheMarinePark,suggeststhatwhilemanagementeffortshavefailedtoallowfishtoincreaseinabundance,theyhavebeeneffectiveatprotectingcertainspecies.Thetsunamihadnodetectableaffectonreeffishassemblagesatthesesites.Thislackofmajordamagemeansthatneithertheconservationprioritiesnortheriskstoreefshavebeenchangedbythetsunamianditisvitallyimportantthatresourcesarenotdirectedtoshortterm,smallscale,rehabilitationprogramswhichwillnotreverselongtermdeclinesinreefconditionwhichwereevidentatmanyofoursites.

INTRODUCTION

Disturbancehasasignificantroleindeterminingthestructureanddynamicsofecologicalcommunities(PickettandWhite,1985;Petraitisetal.,1989),especiallyincoastalmarinehabitats,whichappearparticularlysusceptibletoawiderangeofnaturalandanthropogenicdisturbances(e.g.,Alongi,2002;Hughesetal.,2003).Thesedisturbances,includingseveretropicalstorms,temperaturefluctuations,terrestrialrun-off,anddiseases,varyintheirscale,intensityandfrequency(HughesandConnell1999),contributingtoextremespatialandtemporalvariabilityinthebiologicalstructureofshallow-watermarinecommunities(KarlsonandHurd,1993).Thereisincreasingevidence,however,thateffectsofnaturaldisturbancesarebeingfurthercompoundedbyanthropogenicstressesleadingtodirectionalchangesinthestructureofmarinehabitats.Intheextreme,synergisticeffectsofmultiplechronicdisturbancesleadtoirreversibleandfundamentalshiftsinbiologicalstructure.Oncoralreefs,chronicover-fishingcombinedwithexcessnutrientshasledtopermanentshiftsfromcoral-dominatedtoalgal-dominatedbenthos(Done,1992;Hughes,1994;McCook,1999).Thisinturnmayhavesignificantrepercussionsforthelong-termsurvivalofcoralassociatedreeffishes(reviewedbyWilsonetal.,2006).

CoastalmarinehabitatsinIndonesiahavebeensubjecttoalong-historyofdisturbancefromdestructivefishingpractices(Edingeretal.,1998)combinedwithsevereepisodesofsedimentationandincreasedturbidityassociatedwithmonsoonalrainsandlandbasedrunoff(McManus,1988;HopleyandSuharsono,2002).OnDecember26th,2004,thesehabitatswerefurthersubjecttoanextremepunctuateddisturbanceintheformoftheSumatra-Andamanearthquakeandsubsequenttsunami.Thespatialscaleandmagnitudeofthistsunamihasnohistoricalprecedentandmanyaspectsoftheevent,suchasthelengthofthefaultlineandthespeedoftheslipsuggesteditwasalmostunique(Layetal.,2005,Vignyetal.2005).Estimatesofthereturntimefortsunamisgreaterthan10mwaveheightare1000yearsfortheIndianOcean(TsunamiRisksProject,


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2005)indicatingthatthiswasindeedararenaturaldisturbance.Whilesmallertsunamisarerelativelycommon,forexamplesince1883,35tsunamishaveoccurredinIndonesiaalone(Birowoetal.,1983),thereareasyetfewquantitativestudiesofthedamagetheycausetocoralreefcommunities(Tomascik,1997a,572-4)andconsequentlytheeventprovidedauniqueopportunitytoassesstheeffectsofthistypeofnaturaldisturbanceintropicalmarineecosystems.

InitialreportsofdamagetocoralreefsfollowingthetsunamisuggestedthatgreatestimpactswereinIndonesiaandtheAndamanIslands(UNEP,2005).InIndonesia,initialassessmentsbasedonsatelliteimagerysuggestedthat97,250haofcoralreefhabitatwasaffectedwithapotentiallossof3061havaluedat$332milliondollars(Anon,2005).Regionreportshavesincerevealedthattsunamidamagevariedwidely,andoftenunpredictably.Forexample,Bairdetal.(2005)describedthedamageasoccasionallyspectacular,butsurprisinglylimited,giventheproximityoftheirsitesinAcehtotheepicentreoftheDecember26,2004earthquake.DamagetothereefsofThailand(Comleyetal.,2005;PhongsuwanandBrown,2007)andtheMaldives(Gunnetal.,2005)wassimilarlypatchy,butgenerallylow.Incontrast,widespreaddamagewasreportedtoreefhabitatsintheAndamanandNicobarislands(Kulkarni,2001),SriLanka(CORDIO,2005a;MeynellandRust,2005)andeventheSeychelles(OburaandAbdulla,2005),whichisperhapssurprisinggiventhedistancefromtheepicenteroftheearthquake.TheonlystudytopresentdatafrombothbeforeandafterthetsunamidetectednochangetoshallowcoralassemblagesonPulauWehinAceh(Bairdetal.,2005),despiteanestimatedrun-upheightof5matthislocation(USGS,2005).

InthisstudyweassessedtheconditionofcoralreefsinnorthernAcehregionofSumatratodeterminetheeffectoftheSumatra-Andamanearthquakeandtsunamioncoralreefcommunities.Thestatusofcoralreefcommunities(bothcoralandfishcommunities)wasexaminedagainstabackgroundofconsiderablepriordisturbance.Mostimportantly,reefsinnorthernAcehhavebeensubjecttodestructivefishingpractices,suchascyanidefishingandbombing,whichhavedevastatingeffectsonfishstocksaswellasthebenthicreefhabitats.Accordingly,wesampledsitesunder3differentmanagementregimes;openaccessareas,PulauRubiahMarineReserve,andthetraditionAcehnesemanagementpractice,PanglimaLaut.

METHODS

InApril2005(<100daysafterthetsunami)wevisited49sitesinnorthernAcehlocatedwithin300kmoftheepicentreoftheearthquake(Fig.1).Studysiteswerelocatedwithinthreedifferentmanagementregimes;1)acentralgovernmentmanagedmarinetourismreservecenteredaroundPulauRubiah,whichwewillcallKawasanWisata,2)communitybasedtraditionalAcehnesemarinemanagementsystemknownasPanglimaLaut,and3)openaccessareas.TodocumentcurrentreefconditionandassesspotentialtsunamidamageweusedtherapidassessmenttechniquesrecommendedbytheWorldConservationMonitoringCentre(CORDIO,2005b).Reeffishabundanceanddiversitywerealsoassessedasubsetofthesesites.


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Figure 1.Locationofsitesforassessmentofcoralreefsubstratevariables(47sites)andcoralreeffish(31sites),northernAceh,Indonesia.


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Surveyswereconductedat47ofthe49sitestoassessthebiologicalandphysicalstructureofthereefbenthos,andalsotoquantifyrecentphysicaldamageattributabletothetsunami(Fig.1).Ateachsite16-32replicate10x1mbelttransectswereconductedonthereefcrest(0-2m)and/orthereefslope(3-10m).Onthesetransectsthepercentagecoverofthefollowingvariableswasrecorded,threedescribingreefcondition:1)livecoralcover,2)coralrubble,and3)turfalgae;andthreeindicativeofrecentreefdamage:1)coralcoloniesthatwereup-turnedordisplaced(Fig.2E),2)attachedcolonieswithpartialmortalityorbrokenbranches(Fig.2F),3)recentlykilledcolonies(Fig.2B).ThefollowingcategorieswererecordedasestimatesofcoverfollowingCORDIO(2005b):0%=0;1-10%=5;11-30%=20;21-50%=30;51-75%=62.5;76-100%=87.5).Forstatisticalanalysis,themid-pointofeachcategorywasusedtocalculatemeanvaluesforeachgroup.

Toassesspotentialimpactsofthetsunamionreefassociatedfauna,speciesdiversityofreeffishassemblageswasquantifiedduring20mintimedswimsat31sites.Twodivers(SP,TK)swamalongapre-designatedpathrecordingallspeciesobservedandthelistscombinedtoprovideanestimateofspeciesrichnessforeachsite.Surveyswereconductedalongazig-zagpathstartingat~25mdepthandextendingtothereefcrest.Thetotalareasurveyedwasapproximately300mx100mpersite.

Theabundanceoffisheswithineachof45majorreeffishfamilieswasdocumentedat13sites:3locatedwithinKawasanWisatawhereallfishingisprohibited;3withinPanglimaLautwhereonlyartisanallinefishingispermitted;theremaining7siteswerelocatedinopenuseareas,wherefishingactivitiesarelargelyunregulated,andincludesline-fishing,muro-ami(aparticularlydestructiveformofnetting),netting,trapping,andspearfishing.Thesizeandnumberofallfisheswithineachof45familieswererecordedsimultaneouslyusing3replicate50mtransectsonthereefcrest(<2m).Transectswererunparalleltothereefcrestandspaced>5mapart.Thetransectlinewasdelineatedusinga50mfibreglasstape,alongwhichsmallfishes(<10cmTL)weresurveyedina2mwidepathandlargerfishes(>10cmTL)weresurveyedina5mwidepath.

Thedifferentregimesunderwhichsitesweremanagedshouldinfluencereefcondition.Consequently,wetestedforsignificantdifferenceinmeancoverofcoral,filamentousalgaeandcoralrubbleamongmanagementzonesusinga2-wayANOVA.Factorsinthemodelweremanagement(fixed;3levels,asdescribedabove)andsitenestedwithmanagement(random;4to28sitespermanagementregime).Forthesevariablestheanalysiswasrepeatedtwice;onceforshallowsites(n=38),andagainfordeepsites(n=45)becauseatmanysitestransectswereonlyrunatonedepth.

Tsunamirun-up,whichwasevidentthroughouttheregionasaprominentscarfromwhichvegetationhadbeenstripped,washigherinPulauAcehandthemainlandwhencomparedtoPulauWeh.MeasurementsbytheUnitedStatesGeologicalSurvey(USGS)confirmedtheseobservations,recordingmaximumrun-upheightsinPulauAcehandthemainlandas22mand26mrespectively,4to5timeshigherthanonPulauWeh(~5m)(USGS,2005).Inaddition,ourinitialobservations(seeBairdetal.,2005)suggestedthatdamagewashabitatspecific,inparticular,up-turnedcoralsappearedtobemoreabundantatdepth(>2m)thanintheshallows(<2m).Consequently,weuseda3way-ANOVAtotestformeandifferencesintheproportionofthe3damagevariables(up-turnedcoral,brokencoral,recentlykilledcoral)amonglocations,sitesandbetween


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A B

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G H


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depths.Factorsinthemodelwerelocation(fixed;2levels,PulauAceh/mainland,PulauWeh),sitenestedwithinlocation(random;17and24levels)anddepth(fixed:2levels,shallowanddeep)whichwascrossedwithbothlocationandsitenestedwithinlocation.Onlyup-turnedcoraldifferedsignificantlybetweendepths,sotoincreasethequantityofdatafortheothervariableswerana2-wayANOVAasdescribedaboveforthereefconditionvariableusingtransectsfromeachdepth.

Muchworkontsunamidamagetocoastlinesindicatesthattheangleofincidencebetweenthetsunamiandthecoastlinecaninfluencethedegreeofdamage,andshorelinesfrontingthetsunamiwouldbeexpectedtosuffergreaterdamagethanshorelinesintheleeofthetsunami.Consequently,weuseda2-wayANOVAtotestfordifferencesinthemeanproportionofup-turnedcoral,brokenandrecentlykilledcoralamongsitesfacingnorth,south,eastandwest.Factorsinthemodelwerereefaspect(fixed;4levels,north,south,east,andwestfacingreefs),andsitenestedwithinreefaspect(random;3to19siteperaspect).Onceagain,toincreasethedataavailableforanalysisshallowanddeeptransectswereanalysedseparately.Alldamageandreefconditionvariableswerearcsinetransformedandthenormalityandhomoscedasticityofthetransformeddataexaminedwithgraphicalanalysesoftheresiduals.AnalyseswerecompletedusingSYSTATv10.2.

Coralsreeffishmayalsobeinfluencedbydifferentfishingrestrictionsenforcedwithinmanagementzones.Consequently,wetestedforsignificantdifferenceinmeanabundanceofcoralreeffishamongmanagementzonesusinga2-wayANOVA.Factorsinthemodelweremanagement(fixed;3levels,asdescribedabove)andsitenestedwithmanagement(random;3to6sitespermanagementzone).Onlyasingleestimateofdiversitywasmadeateachof31sites.Consequently,1-wayANOVAwasusedtotestfordifferencesinmeanspeciesrichnessamongmanagementzones(fixed;3levels)withsitevaluesprovidingthereplicationwithinmanagement.Bothvariableswereloge(x+1)transformedtoimprovehomogeneityandnormality,andanalyseswerecompletedusingSYSTATv10.2.

Toexplorespatialvariationinthecompositionofreeffishassemblages,MANOVAwasusedtotestforvariationintherelativeabundanceoffivemajorfamilies(Acanthuridae,Chaetodontidae,Labridae,Scaridae,SerranidaeandPomacentridae)among13sitesforwhichthesedatawereavailable.Alldatawerelogetransformedpriortoanalysestoimprovehomogeneityandnormality,andanalyseswerecompletedusingSPSSv11.0.

Figure 2.A.HealthycolonyofAcropora muricatain1matsite49,November2000.B.ThesamecolonyasinFig2AinApril2005.Despiteanestimatedwaveheightofover12m,thecolonyisstillintact,however,thetissuehasbeensmotheredbysedimentstirredupbythetsunami.C.HealthyreefintheshallowsofPulauRubiahMarineParksite46inApril2005.D.AcollapsedcolonyofHelioporasp.Site11E.AburiedPoritescolonyinapproximately3mdepth.Interestingly,thiscolonywaslessthan20mfromthehealthyreefinFig.2C,demonstratingthedifferentimpactofthetsunamioncoralsfirmlyattachedtoreeforrockwhencomparedtocoralsgrowinginsandorrubble.F.BrokenbranchesinanAcroporasp.site26in0.5mdepth.Thewoundshavehealed,however,thepolypshaveyettobegingrowingagain,suggestingtheinjuryisrecent,andmostprobablycausebydebrismobilizedbythetsunami.G. AlargePoritescolony,approximately3mdiameterliesburiedonthebeachonPulauBeras,site36.H.AbleachedFavitescolonyatsite27.Theturbidityatsomesites,inparticularonthemainlandandinPulauAceh,wasveryhigh,andcontinuestoposeathreattocoralassemblages.

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RESULTS

AtthesesitesonthenorthandwestcoastofAceh,wherethetsunamiwasmostferocious,theinitialdamagetocoralreefs,whileoccasionallyspectacular(Fig.2G),wassurprisinglylimited.Furthermore,damagewasverypatchywithoftenpronounceddifferencebetweenadjacentsites.Tsunamidamagewaslargelyunpredictable:neitherreefaspect,geographiclocation(aproxyfortsunamiintensity)normanagementzonehadasignificanteffectontheamountofdamage.Theonlyclearpatternswereahigherproportionofup-turnedcoralsatdepthandahigherproportionofbrokencoralsonreefcrestsatPulauAcehandmainlandsites.Reefcondition,however,variedwidelywithintheregionandwasclearlycorrelatedwithmanagementregimes.Coralcoverwashigh,andthecoverofalgaeandrubblelowatKawasanWisataandPanglimaLautsites.Incontrast,coralcoverwaslowandthecoverofalgaeandrubblewashighatopenaccesssites.

Themeanproportionofoverturnedcoralswassignificantlyhigheratdepth(shallowsites:3.3±0.35;deepsites;7.6±0.43;F1,33=9.4,P=0.004).Thispatternwasevidentatmostsites,exceptwherethedamagewaslow,suchasmostPanglimaLautsites(Fig.3A),andatthesesites,notsurprisingly,therewasnodifferenceinthemeanproportionofup-turnedcoralbetweendepths,causinganinteractionbetweendepthandsite(management)(F33,1464=4.1,P<0.001).Whiletherewasconsiderablevariationamongsites(management),themeanproportionofoverturnedcoralsdidnotdifferamongmanagementzones(F2,33=0.500,P=0.611).Allmanagementregimeshadsomesiteswithmoderateabundanceofoverturnedcoralandsomesiteswithnooverturnedcorals(Fig.3A).Neitherreeforientation,norgeographiclocationhadanysignificanteffectontheabundanceofup-turnedcoralsoneitherthereefcrestorreefslope.

ThemeanproportionofbrokenlivecoralwassignificantlyhigherintheshallowsatPulauAcehandmainlandareas(21.6±1.65SE)comparedwithPulauWeh(5.7±0.72SE)(F1,34=6.565,P=0.0145)(Fig.3B)butthispatternwasnotrepeatedatdepth(F3,41=2.3,P=0.09).Theabundanceofbrokenlivecoralwasnotsignificantlyaffectedbymanagement,depth,orientation,orgeographiclocationoneitherthereefslope,orthereefcrest.Theabundanceofrecentlykilledcoralswassimilarlyunpredictable,withafewsiteswithineachlocationexperiencinghighmortality,butatmostsitesnorecentlykilledcoralswhererecorded(Fig.3C).

Damagevariableswerepoorlycorrelated.Transectswithahighproportionup-turnedcoralsdidnot,generally,haveahighproportionofbrokencoral(r2=0.087),orrecentlykilledcoral(r2=0.003).Whiletherewasweakcorrelationbetweenbrokencoralandrecentlykilledcoral,only15%ofthevariationwasexplainedbytherelationship.

Allmeasuresofreefcondition(i.e.livecoralcover,turfalgae,coralrubble)variedamongmanagementzones.CoralcoverwassignificantlyhigherintheshallowsatKawasanWisata(31.7±2.8)andPanglimaLaut(52.2±2.2SE)siteswhencomparedwithopenaccesssites(19.3±0.9)(F2,35,=8.4,P<0.001)(Fig.4A).ThispatternwasevenmorepronouncedatdepthwherecoralcoveratPanglimaLaut(44.8±2.7SE)andKawasanWisata(25.8±1.5SE)siteswas3to10timeshigherthanatopenaccesszones(3.8±0.5)(F2,42=5.4,P<0.008).Incontrast,tothispatternbothturfalgae(F2,35,=8.4,P<0.019;Fig.4B)andrubble(F2,35,=3.7,P<0.035;Fig.4C)were10–20timeshigher


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atopenaccesssites(algae=33.9±1.4SE;rubble=20.4±1.2SE)whencomparedtoPanglimaLaut(algae=17.7±2.4SE;rubble=1.5±0.5SE)andKawasanWisatasites(algae=3±0.9SE;rubble=0.4±0.2SE).

Whilethedirecteffectsofthetsunamionthefunctionofcoralreefecosystemswererelativelyminor,changesinthesedimentregimefollowingthetsunamihavecausedlocalizedmortalityandcontinuetothreatensomereefs.Forexample,apreviouslyflourishingAcroporaassemblageatthesouthernedgeofthefringingreefatLampuuk(site49,Fig.1)wassmotheredbysedimentscausingcompletemortality(Fig.2B)comparedwithprevioussurveysinMarch2003(Fig.2A).WhilethesedeadcolonieswerestillintactinApril2005,byDecember2005theyhadcompletelydisappeared.OtherexamplesofindirecteffectsfromthetsunamiincludebleachedAcroporaandfaviidcolonies(Fig.2H)atsites25,27and28.

Atotalof358speciesofreeffisheswererecordedacrossall28studysitessurveyedduringthisstudy.ThemostspeciosefamilieswerethePomacentridae(59 species),Labridae(47 species),Chaetodontidae(32 species),Acanthuridae(28 species)andScaridae(24 species).Speciesrichnessofreeffishesvariedgreatlyamongsamplesites,rangingfrom14speciesatPulauRusa2(site27,Fig.1)to103speciesatGugob1(site38)onthenorth-eastsideofPalauBeras(Fig.5).Thespeciesrichnessofcoralreeffishesvariedgreatlyevenamongcloselypositionedsites.Forexample,73speciesofreeffisheswererecordedatPaloh(site33)onthesouthernsideofPalauBeras,whereasonly19specieswererecordedatLhoh(site32),located<5kmaway.Meanspeciesrichnessdidnotvaryamongmanagementzonesandrangedfrom36.00±3.46SEatPanglimaLautsitesto48.7±5.47SEatopenaccesssites(F2,28=0.45,P=0.645).

Themeanabundanceofreeffishes(averagedacrossallfamilies)variedbyanorderofmagnitudeamongsites,rangingfrom4900(±167.73SE)fishesperhectareatAnoiHitam1(site41),upto94,968(±68,695SE)fishesperhectareatRubiahChannel(site46)(Fig.6).Theoverallabundanceoffishesvariedgreatlyamongsites(df2,10,F=4.32,P<0.05),buttherewasnosignificantvariationattributabletodifferencesinmanagement(df2,10,F=0.36,P>0.05).ThemostabundantfamilyoffisheswasthePomacentridae,whichaccountedformorethan55.9%ofallfishescounted.ThenextmostabundantfamiliesoffishesweretheAcanthuridae,SerranidaeandChaetodontidae,althoughfamiliescomprisingmostlysmallorcrypticfishes(e.g.,ApogonidaeorBlennidae),whichcompriseasignificantcomponentoftheichthyofaunaoncoralreefs(MundayandJones1998)werenotsurveyed.

Whiletherewaslittledifferenceineithertheabundanceordiversityoffishesamongmanagementzones,thestructureofcoralreeffishassemblagesdidvarysignificantlyamongbothmanagementzones(MANOVA,Pillia’sTrace=1.04,F14,42=3.25,P=0.002)andsiteswithineachmanagementzone(MANOVA,Pillia’sTrace=3.10,F70,182=2.06,P<0.001).ThestructureofcoralreeffishassemblagesatsiteswithintheKawasanWisatawasfairlydistinctive,characterizedbyhighabundanceofAcanthuridae(Fig.7).Similarly,thethreesitesfromthePanglimaLautallhadverysimilarfishassemblages,withmuchhigherabundanceofLabridaecomparedtotheKawasanWisata(Fig.7).Notably,fishesfromthefamiliesAcanthuridae,LabridaeChaetodontidae,andSerranidaealltendedtobemoreabundantatKawasanWisataandPanglimaLautsitescomparedtoopenaccessareas(Fig.7).Variationamongsiteswithin


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Figure 3.Spatialandhabitatvariationindamagevariablesat47sitesinnorthernAceh.Valuesarethemean+onestandarderror.Blackbarsrepresenttransectsrunintheshallows(<2m)andwhitebarsrepresenttransectsrunatdepth(>2m).A.Up-turnedcoral.B.Brokenlivecoral.CRecentlykilledcoral.


66

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Figure 4.Spatialandhabitatvariationinreefconditionvariablesat47sitesinnorthernAceh.Valuesarethemean+onestandarderror.Blackbarsrepresenttransectsrunintheshallows(<2m)andwhitebarsrepresenttransectsrunatdepth(>2m).A.Livecoral.B.Filamentousalgae.C.Rubble.


68

eachmanagementregimewashighestamongopenaccesssites,whichdidnotappeartobegroupedbygeographicproximity.Forexample,LhokWeng1(site11)andGapang(site13),whichareopenaccesssiteslocatedwithin1kmofeachotheronthenorthernsideofPulauWeh,hadverydifferentfishassemblages(Fig.7).ThefishassemblageatGapang,andalsoBateeMeuronon(site3),weremostsimilartothoseofsiteswithinthePanglimaLaut,withhighabundanceofLabridae,ChaetodontidaeandSerranidae,whereasthesefamiliesoffisheswererareatmostopenaccessareas,especiallyLhokWeng1(site11)andTepinPineung(site43)(Fig.7).

DISCUSSION

Ourdetailed,largescaleandquantitativesurveyofthereefsinnorthernAcehclearlydemonstratesthatthefirstreportsoftsunamidamagefromthisregionweregrosslyexaggerated.Thevalueofsuchqualitativeassessmentsmustbequestioned,theyarealltooeasytomake,andbecausetheyaretypicallythefirstavailablenews,theycaptureundueattention.Furthermore,theuncriticalrepetitionofthesestudies(e.g.,Tunetal.2005)mustalsobequestioned,becauseitonlyservestoperpetuatedthemyth,andobscureitsprovenance.TheoverwhelmingpicturefromthemajorityofreportsfromtheIndianOcean(Bairdetal.,2005;Brown,2005;PhongsuwanandBrown,2007;Comleyetal.,2005;Gunnetal.,2005)isthatthedamagecausedtocoralreefsbytheDec26earthquakeandtsunamiwasrarelyofecologicalsignificance,andatoursitesinnorthernAceh,tsunamidamagewastrivialwhencomparedwiththatcausedfromchronichumanmisuse.

Fewclearpatternswereevidentinthetsunamidamageobserved:neitherreefaspect,geographiclocation(i.e.tsunamiintensity)normanagementzone(i.e.reefquality)significantlyaffectedanyofthedamagevariables,withtheoneexceptionbeinghighabundanceofbrokenlivecoralonmainlandandPulauWehreefcrests.Thisisperhapssurprising,andcontrastswithresultsreportedelsewhere(Bairdetal.,2005;Brown,2005;ChatenouxandPeduzzi,2005).However,tsunamisinteractwithsubmarineandcoastaltopographyincomplexwaysandinterference,resonance,andreflectioncanconcentratetheforceofthetsunamiinunexpectedlocations,suchastheleeofislands,smallembaymentsandchannels(TsunamiRisksProject,2005).Theearthquakeof26December2004generatedatsunamiinAcehwhichconsistedofatleast3mainwaves(awavetrain),precededbyaninitialdrawdown(Layetal.,2005).Thefirstwavewasestimatedat12mbyeyewitnessesbeforeitbrokeonthereefsontheAcehnesecoast.Thesecondwavewasconsiderablylarger,withflowheightsatthecoastrangingfrom10.0to15.0m(Borrero2005).Indeed,thenortherntipofAcehandtheislandstothenorthwereineffecthitbytwowavetrains,onefromthenorthandonefromthewest(Borrero2005).Withsuchacomplextsunamieventupsuchalargescaleinanareawithmanyislandsofcontrastinggeographyuntanglingthefeaturesthatmadeonereefmoresusceptibletodamagethananotherispossiblyintractable.


69

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Figure 6. Reef fish abundance (ind.ha-1) (mean ± SE) within 2 geographic regions and 3 management zones (Open Access, Kawasan Wisata, Panglima Laut) in northern Aceh, Indonesia.

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Figure 5. Reef fish species richness in within 3 geographic regions (Mainland, Pulau Aceh, Pulau Weh) and 3 management zones (Open Access, Kawasan Wisata, Panglima Laut) in northern Aceh, Indonesia.


70

Theoneclearpatternwasahigherabundanceofoverturnedcoloniesgrowinginunconsolidatedsubstratumbelow2m.Coralsfirmlyattachedtosolidsubstratumwerelargelyunaffectedbytheforceofthewavesatallsites:damagetothesecoloniesincludedoccasionalbrokenbranches(Fig.2C),presumablyasaresultofimpactswithmobiledebris,butveryfewcoloniesweredislodged.Incontrast,coralsgrowinginunconsolidatedsubstrata,suchassandorrubble,sufferedmuchgreaterdamage:inthesehabitatsmanycolonieswereoverturned(Fig.2D),buried(Fig.2F),ortransported,oftenoverlargedistances(Fig.2G).Despitethisdamageatdepth,wherecoralassemblageswerehealthypriortothetsunami,coralcoverremainedhigh,andtherewaslittleapparentlossofecologicaldiversityorfunction.

Thistypeofdamageisverydifferenttothatobservedfollowinglargestorms,suchashurricanes.Whilehurricanedamagetoreefsisalsopatchy(Woodleyetal.,1981),itisunusualforshallowreefstoescapedamageoverlargescalesfollowinghurricanes(HughesandConnell,1999).Furthermore,fragilemorphologies,suchasbranchingandtabularcorals,aregenerallydisproportionatelyaffectedwhencomparedtomassivecoloniesfollowinghurricanes.Anumberoffeaturesoftsunamisarerelevantforexplainingthisdifference.Inwindwaves,mostenergyiscontainednearthesurface,andwave-inducedwatermotiondecaysexponentiallywithdepth(Yehetal.,1993).Incontrast,inatsunami,waterisinmotionthroughouttheentirewatercolumn(Yehetal.,1993).Wehypothesisethattheinitialrundownofthetsunami,alongwiththefirstwaveofthetsunamitrain,excavatedunconsolidatedsubstratafromaroundthebasesofunattachedcolonies,makingthemsusceptibletodisplacementwheninundatedbythesubsequentwaves.Thedifferentialdamagetounattachedmassivecoloniesatdepthappearstobeauniquefeatureoftsunamisdisturbanceandexplainsthedominanceofmassivecoloniesintsunamidepositsonland(Bairdetal.,2005).

Aninterestingfeatureofouranalysiswasthattransectswithhighproportionsofup-turnedcoraldidnotnecessarilyhavehighproportionsofbrokenlivecoralorrecentlydeadcoral.Thissuggeststhatthetypeofdamageobservedatasiteisstronglyinfluencedbywhatcoralspeciesarepresent.Forexample,thehigherproportionofbrokencoralsonreefcrestsonPulauAcehandmainlandreefscomparedwithPulauWehwasprobablytheresultofhighcoverofHeliopora(unpublisheddata),whichhasabrittleskeletonpronetobreakagefrommobiledebris.Acroporacolonies,incontrast,didnotappearpronetobreakage,andwereveryrarelyup-turned,consequently,siteswherethesespecieswereabundant,suchasintheshallowonPulauWehhadfewbrokencorals.Similarly,largethicketsofAcroporamuricataalbeitrecentlykilled(Fig.2B),remainedintact,despiteanestimatedflowheightatthecoastofover15m(Borrero,2005)atthissite.Itis,therefore,surprisingthatdamagetoAcroporacolonieswassoprominentintheSeychelles,morethan3000kmfromtheepicenteroftheearthquake,wherethemaximumwaveheightwas1.24m(Haganetal.,2007).

OngoingeffectsoftsunamiinApril2005includedanincreaseinturbidityatmanysiteswheresomeAcroporaandfaviidswerebleached(Fig2H),probablyasaconsequenceofprolongedperiodsoflowlight(Fabricius,2005),becausethereisnoindicationofrecentelevatedseasurfacetemperaturesinthearea(NOAA,2005).


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-4

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Figure 7. Canonical Discriminant Analysis of coral reef fish assemblages on Acehnese reefs in April 2005. Canonical variates 1 and 2 account for 38.5 % and 26 % of the variation in community structure among all sites and emphasize differences among management regimes (Kawasan Wista = dark grey, Panglima Laut = light grey, and open access = white). Numbers on each centroid correspond with site numbers shown on Figure 1. Circles plotted represent 95% confidence limits around the centroids for each site. Vectors are structural coefficients of response variables, indicating the relative abundance of different families of fishes at each site.


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Reefconditionvariedwidelywithintheregionandwasstronglyinfluencedbycontrolsonhumanactivity(i.e.managementzone).ReefconditionwasparticularlypoorinPulauAceh(Fig.4),herelongdeadcoloniesandrubblebedswerecoveredwithathickgrowthoffilamentousalgae:scenestypicalofreefsaffectedbybombingandcyanidefishing(Pet-Soedeetal.,1999).However,evenhere,wherethetsunamiwashighlydestructiveonland,therewaslittleevidenceofrecentcoralmortality(Fig.3C).Themostlikelycauseoflowcoveratopenaccesssitesisdestructivefishingpractices,suchasbombingandcyanidefishing,bothofwhichwereprevalentthroughoutIndonesiaintherecentpast(HopleyandSuharsono,2002)andmanylocalssuggestedthatsedimentrun-offfrominappropriatelyclearedlandmayhavesmotheredsomereefs(e.g.,LhokWeng–site11andLeunBallee–site40).OnPulauAceh,thesepracticeshavecausedaphaseshift(e.g.Hughes,1994)fromcoralstoalgaewhichthetsunamimayhaveexacerbatedwithaninfluxofnutrientsandtheprospectsforrecoveryofthesereefsintheshorttermarenotgood.

GiventheintensityoftheSumatra-Andamantsunami,itisagainsurprisingthattherewasnoclearevidenceofdisturbancetothereeffishassemblages.Tsunamishavethepotentialtoaffectfishesbydisplacingindividualsorwashingthemashore,ashasbeenobservedduringseveretropicalstorms(e.g.,Walsh,1983).LocalvillagersreportedthatmanysmallfisheshadbeenwashedashoreatPalauWehimmediatelyaftertheSumatra-Andamantsunami(Allen,2005).However,itisthedisturbancetobenthicreefhabitats,suchashighcoralmortalityandmajoralterationsinthephysicalandbiologicalstructureofbenthicreefhabitats,whicharemostlikelytohavethegreatestimpactoncoralreeffishes(Wilsonetal.,2006).Declinesintheabundanceoffishesfollowingextensivedepletionofhardcoralarecommon(e.g.,Sanoetal.,1987;JonesandSyms,1998;BoothandBerretta,2002;Munday,2004;Pratchettetal.,2006),thoughtherecanbeasignificanttimelagbetweenthelossofhabitatandareductioninfishnumbers.Forexample,Pratchettetal.(2006)detectednochangeintheabundanceofobligatecorallivorouscheatodontids,despitea90%declineincoralcoverfollowingcoralbleaching,4monthsaftertheevent,whichsuggeststhatcheatodontidsmaytakelongerthanthistostarveorrelocate.Consequently,thelowabundanceofcheatodontidsatopenaccesssitesmayindicatethatthelowcoralcoveratthesesitespredatedthetsunami.Giventhatwedetectednomajorchangeinbenthichabitatsfromthetsunami,asdescribedabove,itis,therefore,alsohighlyunlikelythatreeffisheswereadverselyaffectedbythetsunami.Whilesignificantspatialvariationintheoverallabundanceandspeciesrichnessofcoralreeffishesamongsiteswasapparent,thiswasnotattributabletodifferentialaffectsofthetsunami.Forexample,theoverallabundanceoffisheswasmuchhigheratTeupinPineung(site43),wheredamagetocoralswasmostpronounced,comparedtoAnoiHitam1(site41),wheretherewasverylittledamagetocorals.However,withoutdatafrombeforetheevent,suchconclusionsmustbetreatedcautiously.

Therelativeabundanceofsomecoralreeffishes,especiallytheAcanthuridae,Serranidae,LabridaeandChaetodontidae,washigherwithintheKawasanWisata(whichisclosedtoallbutlinefishing)whencomparedtoopenaccessandPangLimaLautsitessuggestingmanagementhasbeeneffectiveatprotectingsomespecies,inparticular,those


73

oftencaughtwithnets(Russ,2002).However,totalabundanceofreeffishdidnotvarybetweenmanagementzones,andheavilytargetedfishes,suchaslethrinids,wereinlowabundanceatallsites.Clearly,managementoftheKawasanWisatacouldbeimproved,andtherewasoccasionalevidenceofbreachesofregulations,suchasdiscardednets.However,comparisonsamongmanagementzonesareconfoundedbydifferencesintheaspectandbenthichabitatsofregulatedareasversusopenaccessareas.Thetwoexistingregulatedareas,theKawasanWisataandPanglimaLaut,arebothlocatedonthenorth-eastsideofPalauWeh.Inaddition,thereislittletruereefdevelopmentonPulauWeh:intheshallows,coralsgrowattachedtolargerocks;atdepthPoritesbombieswhichcangrowinsandaredominant(unpublisheddata).Incontrast,reefsonPalauAcehandthemainlandaretruefringingreefswithpotentiallygreaterhabitatdiversity.ThismayexplainwhyspeciesrichnessoffisheswithintheKawasanWisataandPanglimaLautwasoftenlowercomparedtoopenaccessareas.Responsesoffishestoprotectionfromfishingareinfluencedbymanycomplexfactors,includingthesizeofreef,thestructureofreeffishpopulations,theproximityofotherreefsandthelevelofcompliancewithprotectionregulations(Babcocketal.,1999;McClanahanandMangi,2000;Jennings,2001;ShearsandBabcock,2003;Cinneretal.2005).Nonetheless,MPAsaregainingincreasingacceptanceamongscientistsasoneofthefeweffectivewaysofmanagingfisheriesofcoralreefspecies(Russ,2002),andmaybecriticalinmakingreefsmoreresilienttoacutenaturalandanthropogenicdisturbances(Bellwoodetal.,2004).

CONCLUSIONS

FewnaturaleventscancompareinscaleandintensitytotheSumatra-Andamantsunami,yetdirectdamageonreefswassurprisinglylimited,andtrivialwhencomparedtotheclearlossofcoralcoverwherehumanaccesshasbeenuncontrolled.Theextentofthedamageonland,andthetragichumancostshouldnotdistractattentionawayfromtheperennialproblemsofmarineresourcemanagementinIndonesia:improvingwaterquality,reducingfishingpressureandsensiblecoastaldevelopment(Bellwoodetal.,2004).Neithertheconservationprioritiesnortheriskstoreefshavebeenchangedbythetsunamianditisvitallyimportantthatresourcesarenotdirectedtoshortterm,smallscalerehabilitationprogramswhichwillnotreverselongtermdeclinesinreefcondition(Hughesetal.,2005).ThepoliticalgoodwillandthefinancialresourcesthetsunamihasgeneratedshouldratherbeusedtobuildsustainableeconomiesandjustsocietiesthatwillprovidelongtermsecurityforthepeopleofAcehandbeyond.

ACKNOWLEDGEMENTS

WethankS.Connolly,T.P.Hughes,A.HelfgottandM.J.Marnaneforcommentsonthemanuscript.NurmaofOongBungalowsprovidedwonderfulsupportinthefield.ThisstudywasfundedbyUSAidandtheAustralianResearchCouncil.


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disturbance to coral reefs in aceh, northern sumatra

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