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Title: Comparison and cost-benefit analysis of PIT tag antennae resighting and seine-net recapture techniques for survival analysis of an estuarine-dependent fish
Series Title: Fisheries Research 121-122: 153-160
Physical Description: Journal Article
Creator: Barbour, Andrew
Publisher: Elsevier
Place of Publication: Fisheries Research
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Abstract: Studies of fish ecology are enhanced by precise and accurate knowledge of survival, which can be estimated from capture-recapture/resighting based survival probabilities. We conducted a cost-benefit analysis of resighting by an array of 11 autonomous PIT tag antennae and recapture by seine netting, and compared the effectiveness of the two methods for recapturing/resighting marked fish in an estuarine environment. During three separate marking periods, we marked a total of 2109 fish with PIT tags, recap- turing 106 by seine (5.0%) and resighting 1700 by antennae (80.6%). Antennae resulted in precise monthly survival estimates while seine netting did not, but antennae did not collect ancillary data (e.g., growth) and their use was limited to areas where fish used constricted passes <10–30 m in width. Despite a reliance on seine nets to capture fish for marking and high initial construction costs, the cost-effectiveness of PIT tag antennae (US$45–$57 per unique fish resighted) exceeded that of seine netting (US$167–$934). Considering physical capture was required to mark fish, the use of PIT tag antennae is a dual-method approach incorporating both physical captures and telemetry. This dual-method approach can collect cost-effective and highly detailed data that could enhance our ability to make informed management and conservation decisions.
Acquisition: Collected for University of Florida's Institutional Repository by the UFIR Self-Submittal tool. Submitted by Andrew Barbour.
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FisheriesResearch121-122(2012)153…160 Contentslistsavailableat SciVerseScienceDirectFisheriesResearch journalhomepage: www.elsevier.com/locate/fishres Comparisonandcost-bene“tanalysisofPITtagantennaeresightingand seine-netrecapturetechniquesforsurvivalanalysisof anestuarine-dependent“shAndrewB.Barboura ,AaronJ.Adamsb,TannerYessc,DonaldC.Behringera,R.KirbyWolfedaProgramofFisheriesandAquaticSciences,SchoolofForestResourcesandConservation,TheUniversityofFlorida,POBox110600,Gainesville,FL32 611,USAbCenterforFisheriesEnhancement,FisheriesHabitatEcologyProgram,MoteMarineLaboratory,CharlotteHarborFieldStation,POBox529,SaintJam esCity,FL33956,USAcDepartmentofBiologicalSciences,NorthernKentuckyUniversity,HighlandHeights,KY41099,USAdHabitatProgram,MarineFisheriesSection,CoastalResourcesDivision,GeorgiaDepartmentofNaturalResources,OneConservationWay,Suite300, Brunswick, GA31520-8687,USA articleinfo Articlehistory: Received17November2011 Receivedinrevisedform10January2012 Accepted16January2012 Keywords: Centropomusundecimalis Mark-recapture Openpopulationsurvival Juvenile“sh Telemetry Marineabstract Studiesof“shecologyareenhancedbypreciseandaccurateknowledgeofsurvival,whichcanbe estimatedfromcapture-recapture/resightingbasedsurvivalprobabilities.Weconductedacost-bene“t analysisofresightingbyanarrayof11autonomousPITtagantennaeandrecapturebyseinenetting,and comparedtheeffectivenessofthetwomethodsforrecapturing/resightingmarked“shinanestuarine environment.Duringthreeseparatemarkingperiods,wemarkedatotalof2109“shwithPITtags,recapturing106byseine(5.0%)andresighting1700byantennae(80.6%).Antennaeresultedinprecisemonthly survivalestimateswhileseinenettingdidnot,butantennaedidnotcollectancillarydata(e.g.,growth)and theirusewaslimitedtoareaswhere“shusedconstrictedpasses<10…30minwidth.Despiteareliance onseinenetstocapture“shformarkingandhighinitialconstructioncosts,thecost-effectivenessof PITtagantennae(US$45…$57perunique“shresighted)exceededthatofseinenetting(US$167…$934). Consideringphysicalcapturewasrequiredtomark“sh,theuseofPITtagantennaeisadual-method approachincorporatingbothphysicalcapturesandtelemetry.Thisdual-methodapproachcancollect cost-effectiveandhighlydetaileddatathatcouldenhanceourabilitytomakeinformedmanagement andconservationdecisions. 2012ElsevierB.V.Allrightsreserved. 1.Introduction Ecologicalstudiesandeffective“sherymanagementstrategiesareenhancedbypreciseandaccurateknowledgeofsurvival, whichisavitalparameterfordeterminingapopulations“tness( Crone,2001 ).Capture-recapture/resighting(CR)methodsare commonlyusedtodeterminesurvivalforamarkedpopulationby collectingdataforcalculationofapparentsurvivalprobabilities ( =1 Š emigration Š mortality)( Pineetal.,2003 ).Mathematical survivalprobabilitiesareeffectivetoolsforexploringpopulation dynamicsandtestingecologicalhypotheses( Pollocketal.,1990 ), butthequalityofCRdatalimitstheprecision,accuracy,andutility ofsurvivalprobabilities. Correspondingauthor.Tel.:+17034070251;fax:+12392832466. E-mailaddresses: snook@u”.edu (A.B.Barbour), aadams@mote.org (A.J.Adams), yesst1@nku.edu (T.Yess), behringer@u”.edu (D.C.Behringer), Kirby.Wolfe@dnr.state.ga.us (R.K.Wolfe).ThequalityofCRdataislimitedbymethodology.Methods allowingbothahighnumberof“shtobemarkedandrecaptured aredesirable,butCRmethodsinestuarineenvironmentstypically resultinahighnumberof“shmarkedandalownumberrecaptured (e.g., Leberetal.,1998;Hampton,2000 ),oralownumbermarked andahighnumberrecaptured(e.g., HeupelandSimpfendorfer, 2002;Adamsetal.,2009 ).Alowrecaptureratereducestheutilityofsurvivalestimates,whereasmarkingtoolowanumberof“sh resultsinanon-representativestudypopulation. ToadvancetheuseofCR-basedsurvivalestimatesincoastal estuarinesystems, Adamsetal.(2006) appliedthe“rstestuarine versionofanautonomouspassiveintegratedtransponder(PIT)tag antenna.ThisCRsystemallowedforahighnumberof“shtobe markedwithlowcostPITtags,andresighted(asindividualswere notphysicallyrecaptured)ahighnumberofmarkedindividuals. Thesingleantennain Adamsetal.(2006) resighted>40%of314 markedindividuals,aresightingraterepeatedby Meyneckeetal. (2008) .Althoughthissingle-antennaapproachprovidedsuperior apparentsurvivalestimatestopreviousapproaches( Adamsetal., 2006 ),precisionwouldbeincreasedandbiasreducedbyusing0165-7836/$…seefrontmatter2012ElsevierB.V.Allrightsreserved. doi: 10.1016/j.“shres.2012.01.013

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154 A.B.Barbouretal./FisheriesResearch121-122(2012)153…160multipleantennaewhenstudyingmobilespecieswithcomplex habitatrequirements. Weexpandeduponthisapplicationwithamulti-antennaarray tostudythejuvenilelife-stageofanestuarine-dependent“sh,the commonsnook Centropomusundecimalis .Weconstructedeleven antennaeoverfourtidal-mangrovecreeks,andcomparedresightingofPITtagsbyantennaetorecapturebyseinenetting.The purposeofthispaperwasstrictlytocompareantennaandphysical CRmethodsusingacost-bene“tframeworksothatotherscanevaluatetheapplicationofthismethodfortheirsystemandresearch goals. 2.Materialandmethods 2.1.Studyarea CharlotteHarborisa700-km2coastalplainestuaryinsouthwestFlorida(USA)( Hammett,1990 ).Theclimateissubtropical, withmeanseasonalwatertemperaturesrangingfrom12Cto36C andinfrequentfreezingairtemperatures( Poulakisetal.,2003 ). Seagrassmeadows(262km2; Sargentetal.,1995 )dominatethe benthichabitatandmangrovesdominatetheshoreline(143km2; Kish,unpublisheddata).Thisstudywasperformedinfourred mangrove( Rhizophoramangle )fringed,tidal,estuarinecreeks,each approximately1.6kmlong,ontheeasternshoreofCharlotteHarbor.Thecreeksvariedinwidthfrom2mpassesto>60mbays,and averagedepthsrangedfrom0.5to2.0m,withthedeepestoccurring inthenarrowpasses. 2.2.Cost-bene“tanalysis Toconductacost-bene“tanalysisofrecapturebyseinenetand resightingbyantennae,we“rstcompiledtheoverallcostsassociatedwitheachmethod.Forseinenetting,thecostsconsistedof theseinenet(US$1000),thepassiveintegratedtransponder(PIT) tagsused(US$2.50tagŠ 1),fuelcosts(US$0.80LŠ 1),andlaborcosts (US$15hŠ 1).Thesumoftheseexpendituresencompassedboth seinenetmarkingandrecapturecosts.Forantennaeresightings, westartedwiththecostofseinenetting…asseinenettingwas requiredtomark“sh…andaddedthecostofconstructionmaterials andlabor(US$15hŠ 1)requiredtobuildantennae. Tocomparethebene“tofeachmethod,we“rstcalculated theperyearcostofeachapproach.Forbothmethods,equipment usedformultipleyears(e.g.,seinenets,antennae)hadinitialcosts spreadoverallyearsused.Next,wedeterminedthecostofrecapturing/resightinguniquelymarkedindividualsbothasingletime, andonamonthlybasiseachyear.Wethencomparedtheutilityof thedatacollectedwitheachmethodbyfocusingonthequalityof apparentsurvivalestimates.Finally,wediscusstheabilityofeach methodtocollectancillarydata(e.g.,growth,populationsize)and commentonthegeneraluseofeachmethod(e.g.,suitabilityfor differenthabitats,utilityindifferentsamplingconditions,etc.)as qualitativemeasuresofbene“t. 2.3.Focalspecies Commonsnook( C.undecimalis )isasubtropical/tropical, estuarine-dependent,euryhalinespeciesthatisecologicallyand economicallyimportantthroughoutitsrange,especiallyinFlorida ( Tayloretal.,2000 ).Adult C.undecimalis spawninpassesandinlets atthemouthsofestuariesinsalinities 25ppt( Tayloretal.,1998 ); thenearshoreplanktoniclarvalstagelastsapproximately2.5wk ( Petersetal.,1998 );andjuvenilessettleintoshallow,mesohalineto oligohalinehabitats( Petersetal.,1998 ).Juvenile C.undecimalis are commoninornearmangrovecreeksyear-round,withhighestdensitiesinthefallandwinter,untiltheyreachapproximately300mm Fig.1. SchematicofPITtagantennaandcomponentparts.Partsinclude:(a)inductor coilinswim-throughorientation;(b)initialinductorcoilcablelength;(c)tunerbox; (d)twinaxialwire;(e)readerbox;(f)batterybox;(g)junctionbox;and(h)solar panel.standardlength(SL)( Tayloretal.,2000;Adamsetal.,2006 ),when theybegintoentertheadultpopulation. C.undecimalis largerthan 300mmSLuseopenestuarineandnearshorehabitats(e.g.,mangroveshorelines,arti“cialstructure)fromspringthroughfall,and presumablyoverwinterinriverineorcreekhabitats( Blewettetal., 2009 ). 2.4.Antennasystems PITtagantennaewereoriginallydesignedforarti“cialfreshwaterenvironments,suchashydroelectricdams(e.g., Castro-Santos etal.,1996;Giorgietal.,1997 ).Recently,twogroupssuccessfully designedsinglePITtagantennasystemsforapplicationinnatural marineandestuarinesystems( Adamsetal.,2006;Meyneckeetal., 2008 ).Weconstructedantennaebasedonadesignsimilarto Adams etal.(2006) .Thesystem( Fig.1 )consistedofanopenloop,copper inductorcoilantenna(asingleloopof660-strand,6gaugecopper weldingcable)connectedtoatuningbox,inturnconnectedtoa readerboxcontainingadata-loggingcomputer(tunerandreader boxespurchasedfromOregonRFID).Eachantennaoperatedcontinuouslywithpowerfromtwo6Vbatteriesconnectedinseries andchargedbya130Wsolarpanel.Thisantennadesignpermittedsubstantiallygreatercoverageofcreekwidththanthe”atplate antennaof Hewittetal.(2010) ,butrequiredamoderatedegreeof self-fabrication. PITtagscontainnobattery,whichallowsforaninde“nitelifespan( GibbonsandAndrews,2004 ),butrequiresthetagtobeinclose proximitytotheantennaforresighting.ToreadaPITtag,electric charge”owsthroughthecopperinductorcoiloftheantenna,producingamagnetic“eld.Whenamarked“shpassestheinductor coil,themagnetic“eldinducesachargeonacoilofwirearounda ferritecoreinthePITtag.Theactivatedtagtransmitsitsuniquely codedIDnumbertotheantennasystem.ThereadrangeŽofan antennaisde“nedasthemaximumdistancefromaninductorcoil thataPITtagcanberead.Acomputersystemstorestheresightingdataandwasretrievedbyconnectingalaptopcomputeror personaldigitalassistant.Thecopperinductorcoilwasoriented horizontally(”atbed: Armstrongetal.,1996 )orvertically(swimthrough: Adamsetal.,2006 )inthewatercolumn,dependingupon location.Weusedthesingleswim-throughdesignantennafrom Adamsetal.(2006) ( Fig.1 ),anddeployedtenadditional”atbed antennae.Flatbedantennaewereidenticaltotheswim-through antenna,excepttheinductorcoil( Fig.1 a)wasrotated90andlaid ”atacrossthebottomofthecreek.Weconstructed”atbedantennaetoavoidentanglementbyrecreationalboatmotorsandbecause

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A.B.Barbouretal./FisheriesResearch121-122(2012)153…160 155 Fig.2. DiagramofthestudycreeksinCharlotteHarbor,FLUSA.Starsrepresent antennaelocations.C.undecimalis tendtoswimlowinthewatercolumn( Petersonetal., 1991 ),increasingtheprobabilityofdetectionbythe”atbeddesign. The”atbeddesigncreatedaPITtagdetection“eldupwardfromthe bottom,whereastheswim-throughdesigndetectedtagswithin theareaencircledbytheinductorcoil.Weplacedoneantenna every0.5kminthelower,middle,andupperstrataofeachcreek withtheexceptionofYuccaPenupper(duetocostconstraints) ( Fig.2 ). Thedimensionsoftheinductorcoillimitedtheapplicationof PITtagantennatoconstrictedstretchesofwater,butinother studiesantennahavebeenconstructedwith30mlonginductorcoils(J.VincentTranquilli,OregonDepartmentofFishand Wildlifepersonalcommunication).Usinga12Vbatterysystem andweldingcablewith660copperstrands,weconstructedrectangular”atbedinductorcoilswithasingleloop,dimensionsof 9.5m 0.75m,andaninitialcablelengthof1.0m( Fig.1 b).For theequipmentandmaterialsused,thesedimensionsmaximized readrange( Barbouretal.,2011 ).Antennacoilswererunthrough mangroveproprootsoneithersideofthecreekorstakedto thegroundusingsectionsofPVCpipetomaintaintheproper dimensions.Totestreadrange,werepeatedlypasseda23mmhalfduplexPITtagovereachantennaatvaryingdistancesuntilthe tagwasnolongerdetected.Duetoavailabilityofsuitablenarrow creekstretches,fewantennaecovered100%ofcreekwidth,butall covered>75%. 2.5.Markingandseinenetrecapture Usingacenterbagseine(30.5 1.8m,6.3mmmesh),juvenile C.undecimalis werecaptured(120…300mmSL)betweenFebruary 2008andFebruary2011inthreeseparatemarkingperiods.For markingperiodone,seine-netsamplingoccurredoveratotalof 44days:FebruaryandMarch2008(1dayeach),April2008(3days), May2008(2),June2008(4),July2008(7),November2008(5), December2008(13),JanuaryandFebruary2009(3dayseach), andAprilandJune2009(1dayeach).Formarkingperiodtwo (9days),wemarked“shinNovember2009,January2010,and February2010(3dayseach).Formarkingperiodthree(14days), wemarked“shinOctober2010(3days),November2010(5), December2010(1),January2011(4),andFebruary2011(1).These physicalcaptureeventswereusedbothtomark“shandforseinenetrecapture.Seine-netsamplingoccurredonlyduringperiodsof lowtide,when C.undecimalis wereunabletoseekrefugeamong mangroveproproots.Seinenettingoccurredthroughoutthelength ofeachcreek.Individualseinenetpullswerenotstandardized, butinsteaddesignedtomaximizecaptureef“ciencyforindividual pulls.Forexample,inSouthSilcoxCreek,weoccasionallytrapped juvenilesinalong,narrowditchusingtwoseinenetspulledinto eachother. Uponseinecapture,wescannedall“sh(documentingphysicalrecaptures)withahandheldPITtagreader(modelno.RS601, All”ex)andmeasuredSLtothenearestmillimeter.Wemarked “shwithuniquelycodedhalf-duplex(HDX)PITtags(23mm length 3.4mmdiameter,0.6ginair;TexasInstrumentsTIRFIDS2000).Weinsertedtagsintotheabdominalcavityofallunmarked “shthrougha3mmincisionposteriorandventraltothepectoral “n.Forthismark,apreviousstudyfound100%tagretentionwith nomortalityforjuvenile C.undecimalis >120mmSL,andnoneed forsuturestoclosetheincision( Adamsetal.,2006 ). 2.6.Survivalmodelselectionandanalysis Wecreatedmonthlycapturehistoriesforantennaebycombiningantennaeresightingdatawithseinenetmarkingevents. Weassignedeachindividuala1Žinmonthsresightedormarked, anda0Žinmonthsnotresightedormarked.Wecollapsedantennaeresightingandseinemarkingdataintomonthlysamplingbins fromFebruary2008toJune2009( n =17)formarkingperiodone, November2009toAugust2010( n =10)formarkingperiodtwo, andOctober2010toJune2011( n =9)formarkingperiodthree. Forseine-netcapturehistoriesinmarkingperiodone,wecollapsedseine-netsamplingeventsintodiscretetimeperiods.We thenscaledthetimeintervalsbetweenseine-neteventssothata 30-dayperiodequaledanintervaloflength1.The n =17seine-net intervallengthswerecodedasfollows:1.03,0.70,1.40,0.43,0.30, 0.97,0.33,4.2,0.13,0.40,0.10,0.27,0.17,0.57,0.90,2.30,and1.50. Wedidnotcreatecapturehistoriesforseinenettinginmarking periodtwoorthreeduetoaninsuf“cientnumberofrecaptures. Followingtheexampleof Adamsetal.(2006) ,weused aCormack…Jolly…Seber(CJS)openpopulationmodel( Cormack, 1964;Jolly,1965;Seber,1965 )inthecomputerprogramMARK ( WhiteandBurnham,1999 )toestimateapparentsurvival.Bycollapsingcontinuous-timeresightingdataintodiscretetimebins forantennaecapturehistories,weviolatedanassumptionofthe CJSmodel(instantaneoussamplingperiods).However,throughan ongoingsimulationstudy,wehavedemonstratedthatthemean biasassociatedwiththisviolationisontheorderof Š 4.0to0.0% (A.B.B.,unpublisheddata)fortheestimatedparametervaluesin thisstudywhenusingone-monthintervals. TheCJSmodelcalculatestwoparameters:(1)apparentsurvivalprobability( =1 Š mortality Š emigration),and(2)capture probability( p ).Forsimplicityofsurvivalanalysisandcomparison

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156 A.B.Barbouretal./FisheriesResearch121-122(2012)153…160betweengeartypes,weeitherestimateduniqueparametervalues onatime-dependent( t )orindependent( )basis.Fixingaparameterintime( )returnsasingleparametervalue,whichrepresents asinglevalue“tbetweenalltimeintervals,asopposedtocalculatinguniqueparametervaluesbetweenalltimeintervals( t ).We builtandcomparedfoursimplemodelsthatallowedeachofthese twoparameterstoeithervaryorbeconstantovertime.Capture probability( p )was“xedat0fortimeintervalsantennaewerenot activeorseinenettingdidnotoccur.Forantennaeresightingsin markingperiodone,parametersestimatesareonlygivenfromJuly 2008toJune2009sinceantennaewerenotconstructeduntilJuly 2008. Toselectthemostappropriateofthefourpossiblemodels,we usedAkaikesInformationCriterion(AIC)values( Akaike,1973 )and relevantbiologicalknowledgeofthesystem( Pineetal.,2003 ), suchasseasonaltrendsin C.undecimalis lifehistory.Forcorrectionofsmallsamplesize,AICcwasused,whichconvergestoAIC athighsamplesizes.Wede“nedmodelswith AICc( AICc=AICc valueofgivenmodelminusminimumAICcofthefourmodelruns) values<2ashavingsubstantialsupport,4 AICc 7ashaving considerablylesssupport,and AICc>10ashavingnosupport ( BurnhamandAnderson,2004 ).Biologicalknowledgeofthesystemsuggeststheappropriatemodelwillallow p and tovary withtime( ( t ) p ( t )),becauseadult C.undecimalis usemangrove creeksduringwinterandopportunisticallycannibalizejuveniles duringthisco-occurrence( AdamsandWolfe,2006 ).Additionally, juvenile C.undecimalis movementandemigrationvaryseasonally, likelyaffectingboth and p ( Stevensetal.,2007 ;A.J.A.,personal observation). 3.Results 3.1.Cost-bene“tanalysis Althoughweonlyconstructedtenantennae,wealsoincluded thecostoftheeleventhantennaremainingfrom Adamsetal. (2006) .The“nalcostforantennamaterialswasapproximately US$4000perantenna(solarpanel,readerandtuner,batteries, wiring,boxes,andwood: Fig.1 ).Anapproximatetotalof800 person…hours(inexperiencedtwo-personteam)wererequiredto design,fabricate,deploy,tune,andtestallantennae.EachpersonhourwasvaluedatUS$15.Therefore,totalconstructionlabor costUS$12,000(800h US$15perhour).Equipmentandlabor costsfortheantennaetotaledUS$56,000(US$12,000forlaborand US$44,000formaterials). ThePITtagantennaearraytotalconstructioncostwasapproximatelyUS$56,000,resultinginaUS$18,667peryearcost(US$1697 perantenna)during3yearsofdatacollection.Inadditiontoconstruction,seinenettingwasrequiredtomark“sh( Tables1and2 ). Therefore,thepricedifferencebetweenantennaeresightingand seinerecapturewastheper-yearcostofantennaeconstruction (US$18,667).Duringthethreeyearsofuseforthisstudy,the costeffectivenessofantennaeresightingwasevidentwhencalculatedbyuniqueindividualoruniquemonthlyrecapture/resighting ( Table3 ).Thecostofuniquemonthlyrecaptures/resightingswas anorderofmagnitudehigherfortheseine-nettechnique.Table1 Cost(US$)formarkingandseinenetrecaptureinmarkingperiodsone,two,and three.One-timecostswerespreadoverallsamplingperiodsandroundedtothe nearestdollar. PeriodonePeriodtwoPeriodthree Labor($15personhŠ 1)$13,200$2700$4200 PITtags($2.50tagŠ 1)$2608$1483$1183 Fuel($0.80LŠ 1)$748$153$238 Seinenetcost($1000total)$333$333$333 Grandtotal$16,889$4669$5954 3.2.Markingperiodone Wemarked1043juvenilesnook( Table4 )during880person-h ofseinenetting.Thesehourswerealsousedforphysicalrecapture, resultingintherecaptureof91uniquelymarked“sh(110total recaptures)foran8.6%overallrecapture.Antennaewerefunctional fromJuly2008toJune2009.Duringthistime,theten”atbedantennaeaveragedaverticalreadrangeof23.5cm 1.49SEwitharange of11…41cmasmeasuredatmultipletemporalpoints(withsalinitiesranging39.6…6.9 ‹ )…anegativecorrelationexistsbetween readrangeandsalinity( Barbouretal.,2011 ).Antennaeresighted 744uniquelymarked“shatleastonce(>200,000totalresightings) fora71.3%overallresighting( Table5 ).Inmarkingperiodone, antennaeoutperformedseinenettingintermsofoverallrecapture/resightingrate,whilealsoresultinginalmost2000timesas manytotaldetections. ComparingCJSmaximumlikelihoodestimatesofapparentsurvival( )andcaptureprobability( p )fromthe ( ) p ( ) model:antenna =0.767(95%CI=0.751…0.781)and p =0.723 (0.701…0.744),whileseine =0.561(0.471…0.648)and p =0.0427 (0.0329…0.0552).Basedon95%con“denceintervals,bothantennaeapparentsurvivalandcaptureprobabilitiesweresigni“cantly ( =0.05)higherthanseine-netestimates.Additionally,95%con“denceintervalsweresmallerforantennaeparameters,resultingin moreinformativemaximumlikelihoodestimates. AICcresultsfromCJSopenpopulationmodels( Table6 aand b)stronglysupportedthebiologicallyreasonablemodel ( t ) p ( t ) ( AICc=0)fordatafromthePITtagantennaarray,withthenext mostlikelymodelreceivinga AICc=99.4.Datafromseine-net recapturesresultedinsimilarsupportforthebiologicallyreasonablemodel( AICc=0.0)andmodel ( ) p ( t )( AICc=0.74). Therefore,weperformedmodelaveragingonthesemodels( Fig.3 ) baseduponAICcweight( BurnhamandAnderson,2002 )( Table6 b). Examiningmodel-averagedresults( Fig.3 ),therewashigher precisioninantennaethanseine-netCRdata.Forseine-netparameterestimatesbeforemonthlybin5,95%CIswere0.01…0.83or 0.16…0.99forall estimates,and0.0…1.0for p …theseestimates areexcludedfrom Fig.3 .Thehighdegreeofprecisioninantennae andhigh p ( Fig.3 )forthe“nalsixparameterestimatescoincidedwiththefullfunctioningofallantennae,andwiththemarkingof ahighnumberof“sh( Table4 ).Antennaeresightedindividual“sh inmultiplemonthlybinsatasubstantiallyhigherratethanseine netting( Table5 ),drivingprecisemonthlyparameterestimates.For example,tenunique“shwererecapturedintwodistinctmonthsby seinenetting,andno“shwererecapturedinthreeormoremonths.Table2 Cost(US$)formarkingandantennaeresightinginmarkingperiodsone,two,andthree.One-timecostswerespreadoverallsamplingperiodsandrounde dtothenearest dollar. PeriodonePeriodtwoPeriodthree Markingcost(from Table1 )$16,889$4669$5954 Antennaematerials($4000antennaŠ 1 11antennae=$44,000total)$14,667$14,667$14,667 Antennaeconstructionlabor(800h $15hŠ 1=$12,000total)$4000$4000$4000 Grandtotal $35,556$23,336$24,621

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A.B.Barbouretal./FisheriesResearch121-122(2012)153…160 157 Table3 Formarkingperiodsone,two,andthree,thetotalcost(US$)ofeachrecapture/resightingmethod,thecostperunique“shrecaptured/resighted,and thecostperunique monthlyrecapture/resightingroundedtothenearestdollar. PeriodonePeriodtwoPeriodthree SeineAntennaeSeineAntennaeSeineAntennae Totalcost( Table1 ,2)$16,889$35,556$4669$23,336$5954$24,621 Costperunique“shrecaptured/resighted( Table5 )$186$48$934$45$595$57 Costperuniquemonthlyrecapture/resighting$167$14$934$15$541$15 Table4 Cumulativenumberof“shmarkedandnumberrecapturedbyeachgeartypepermonthlybininmarkingperiodone.N/Adesignationrepresentsmonthswhenn ophysical samplingoccurred,ormonthswhenantennaehadyettobeconstructed.Captureprobability( p )was“xedat0forthesemonths. MonthlybinMonthCumulativenumberofmarked“shRecapturedbyseineResightedbyantennae 1February2008220N/A 2March2008280N/A 3April2008970N/A 4May20081170N/A 5June20081300N/A 6July2008212143 7August2008212N/A29 8September2008212N/A31 9October2008212N/A43 10November20085156211 11December200876945354 12January200995534421 13February2009103612384 14March20091036N/A407 15April200910423321 16May20091042N/A178 17June20091043088 Incomparison,515unique“shwereresightedinatleast2months bytheantennaarray,and432inatleast3months( Table5 ). 3.3.Markingperiodstwoandthree Wemarked593juvenilesnookbetweenNovember2009and February2010(markingperiodtwo)during180person-hofseine netting.The180hofphysicalcaptureresultedintherecapture of“veuniquelymarked“sh(5totalrecaptures)fora0.8%overallrecapture.Wemarked473juvenilesbetweenOctober2010and February2011(markingperiodthree)during280person-hofseine netting.Seinenettingresultedintherecaptureoftenuniquely marked“sh(eleventotalrecaptures)fora2.1%overallrecapture. Lowphysicalrecaptureratesresultedininsuf“cientdataforcalculationofmaximumlikelihoodsurvivalestimatesformarking periodtwoorthree.Therefore,thesemarkingperiodsdidnotprovideameaningfulcomparisonofantennaeandseinenetsurvival estimates.Instead,theyhighlightthequalityofsurvivaldatathat canbecollectedbyantennaewithonlyaminimumoftimespent marking“sh(9and14days,respectively). FromNovember2009toAugust2010(markingperiodtwo),the antennaeresighted523( Table5 )uniquelymarked“sh(>100,000 totalresightings)resultingin88.2%ofmarked“shbeingresighted. FromOctober2010toJune2011(markingperiodthree),theantennaeresighted433( Table5 )uniquelymarked“sh(>190,000total resightings)resultingina91.5%overallresighting.Thisresighting dataresultedinstrongAICc( Table6 candd)supportforthebiologicallyreasonablemodel ( t ) p ( t )inbothmarkingperiods.Antennae parameterestimateswerehighlyprecise( Fig.4 ),andasmarked individualswerelostthroughmortalityandemigration,variability inparameterestimatesincreased.Parameterestimatesfrommarkingperiodtwoidenti“edadecreaseinapparentsurvivalbetween thethirdandfourthmonthlybins( Fig.4 ).Thisdecreaseinapparent survivalcoincidedwithaperiodwhentemperaturesfellbelowthe lethaltoleranceofthestudyspecies( Adamsetal.,2012 ). 4.Discussion Forcoastal“shstudieslastingmultipleyears,applicationofthe PITtagantennaemethodisacost-effectivewaytogreatlyincreaseTable5 Incidenceofsingleandmultiplerecaptures/resightingsbygeartype.Recaptures/resightingsrepresentthenumberofunique“shrecaptured/resi ghtedinatleastthegiven numberofmonthlybins.Forexample,inmarkingperiod(MP)onetherewasone“shthattheantennaarrayresightedintwelveseparatemonthlybins.1043 “shwere markedinMPone,593inMPtwo,and473inMPthree. NumberofmonthlybinsAntennaeresightings(MPone)Antennaeresightings(MPtwo)Antennaeresightings(MPthree)Seinenetrecaptures(MPone) 174452343391 251237534310 34322782630 43321562040 5237901530 6147591000 76725510 8239310 98430 1040N/A0 113N/AN/A0 121N/AN/AN/A

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158 A.B.Barbouretal./FisheriesResearch121-122(2012)153…160 Fig.3. ApparentsurvivalandcaptureprobabilitiescalculatedfromresightingbyanarrayofelevenautonomousPITtagantennaeversusrecapturebyseinen etting(uneven timeintervals)of1043 Centropomusundecimalis markedduringmarkingperiodone…February2008toMay2009(monthlybins1…16).Parameterscalculatedusing Cormack…Jolly…Sebermodel-averagedresults.Monthlybinswhenseinenettingdidnotoccurorantennaewerenotconstructedhadcaptureprobabilit y“xedto0andare excludedfromthis“gure.Monthlybinsbeforeantennaeconstructionarealsoexcluded.Barsrepresent95%con“denceintervals.precisionofsurvivalestimatesandprovideotherwiseunavailableinformation.Comparedtoseinenetting,antennaeresighted marked“shatahighrate,allowingforinformativemonthlyestimatesofapparentsurvival.Theantennaearraycollectedsurvival informationwithaminimumofphysicalcaptureeffort,andwasTable6 AICcresultsandnumberofestimatedparameters(np)fromCormack…Jolly…Seber RecapturesOnlyModelinprogramMARKforthe:(a)antennaarray(markingperiod one),(b)seinenetrecapture(markingperiodone),(c)antennaarray(markingperiod two),and(d)antennaarray(markingperiodthree).Apparentsurvival( )andcaptureprobability( p )werecalculatedoneitheratime-dependent( t )orindependent ( )basis. ModelnpAICc AICcAICcweight (a) ( t ) p ( t )275455.30.01.0 ( ) p ( t )135554.799.40.0 ( t ) p ( )175634.3179.00.0 ( ) p ( )25865.3410.00.0 (b) ( t ) p ( t )24936.40.00.59 ( ) p ( t )18937.20.70.41 ( t ) p ( )17991.855.40.0 ( ) p ( )2997.661.20.0 (c) ( t ) p ( t )173045.20.01.0 ( t ) p ( )103092.847.60.0 ( ) p ( t )103108.763.50.0 ( ) p ( )23187.2142.00.0 (d) ( t ) p ( t )152579.80.01.0 ( t ) p ( )92591.912.10.0 ( ) p ( t )92621.041.20.0 ( ) p ( )22647.167.40.0 notlimitedbyenvironmentalconditions.However,whileseine nettingwaslimitedbyenvironmentalconditions(e.g.,suitable tides,severeweather),itwasnotreliantonconstrictedbodiesof waterandisthereforeusefulinagreatervarietyofhabitattypes. Also,whileantennaeweremorecost-ef“cientthanphysicalcapture,theywereunabletocollectbioticcovariates(e.g.,“shsize), whichcouldjustifythehighercostofphysicalrecapturesincertain studies. 4.1.Cost-bene“t Despitetherelativelyhighinitialinvestmentinmaterialsand labortoconstructtheantennae,thecosteffectivenessanddata superiorityfromtheantennaapproachvalidatedtheexpense. Physicalcapturewasrequiredtomark“sh,anecessityofany capture-recapture/resighting(CR)study,butantennaeCRwas morecost-ef“cientthanseinenettingevenwhenconsideringthese markingcosts.Thiscosteffectivenessiscompoundedinlong-term researchprograms,asantennaefunctionformultipleyears…the antennafrom Adamsetal.(2006) hasfunctionedcontinuouslyfor sevenyears.Forlong-termresearchprojects,theonlycostsafter antennaeconstructionaremarking,dataupload,batteryreplacementafter4…7years,andinfrequentmaintenanceexpenditures. Ourcost-bene“testimatesforantennaesystemswereconservativesincewedidnotfactorintheadditionalyearsofpotential useoftheantennaearray.Additionally,actualuseofthesesystemsprovidesfurtherinformation,suchasdataonlong-term site“delityandmovementpatterns,providingadditionalbene“ts beyondenhancedapparentsurvivalestimates. Forseine-netrecaptures,thelevelofphysicaleffortrequiredto replicateantennaeresightingwouldin”atecostsbeyondantennaeconstructionlevels,elevatethemagnitudeofpost-release mortality,andinduceatrap-responsebias( Nicholsetal.,1984 ).

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A.B.Barbouretal./FisheriesResearch121-122(2012)153…160 159 Fig.4. MonthlyapparentsurvivalandcaptureprobabilitiescalculatedfromresightingbyanarrayofelevenautonomousPITtagantennaeof593 Centropomusundecimalis markedduringmarkingperiodtwo…November2009toJuly2009(monthlybins1…9),and473 C.undecimalis markedduringmarkingperiodthree…October2010toMay 2011(monthlybins0…7).ParameterswerecalculatedusingCormack…Jolly…Sebermodel-averagedresults.Barsrepresent95%con“denceintervals.Additionally,theunderestimateofapparentsurvivalinthe ( ) p ( ) modelwasindicativeofanegativebiasrelatedtotemporaryemigration(A.B.B.,unpublisheddata),whileasimilarbiaswaslikely dampenedinantennae-basedestimatesbycaptureprobabilities >0.5( Zehfussetal.,1999 ).Thus,weareunawareofanyreasonablephysicalCRmethodthatwouldresultinsuf“cientrecaptures tomatchtheantennaeresults.Thiswasparticularlyimportant inthisstudy,becausethemagnitudeofseine-netrecaptureswas insuf“cienttogenerateinformativemonthlyestimatesofapparent survival.Incomparison,thetemporallydetailedsurvivaldatafrom antennaeresightingwasusefulasitclearlyidenti“edadisturbanceeventbetweenthethirdandfourthmonthlybinsofmarking periodtwo( Adamsetal.,2012 ).Thistemporallydetailedandhighly precisedatacouldbeusedtoidentifyessential“shhabitatsor inform“sheriesmanagementmodels(e.g.,bypredictingyearclass strength).Additionally,althoughnotincludedinthisstudy,wehave successfullyusedantennaeresightingstogenerateinformative monthlysurvivalestimatesbycreekand“shageinanongoinganalysisofjuvenilesurvival(A.B.B.,unpublisheddata).Thistypeand qualityofdatawouldnotbeobtainablewithaseine-netapproach inthisstudysystem. 4.2.Methodologicalcomparison PITtagantennaewereespeciallyusefulinthisstudy,aswesampledinacomplexmangrovehabitatwhereeffectivesamplingwas oftenchallenging(e.g., RobertsonandDuke,1990 ).Forexample, seinenettingwasonlyef“cientduringlowtidewhenmangrove proprootrefugewasunavailable( Thayeretal.,1987;Laegdsgaard andJohnson,2001 ),buttheantennaefunctionedcontinuously andthroughoutthefulltidalcycle.Furthermore,antennaeproved extremelyadvantageousinthesummermonthswhenlowtides occurredatnightandfewerjuvenilesinhabitedthecreeks.These factorsmadesummerseine-netsamplingdif“cultandinef“cient, returninglittledatafortheeffortexpended.Antennaeresighting data,ontheotherhand,allowedsurvivalanalysistocontinueinto thesummerandlentinsightintoseasonalcharacteristicsofhabitatuse,suchasemigrationorontogeneticshiftfromthejuvenile habitat. OnechallengetousingPITtagantennaesystemsinestuarine environmentswasachievingproperdesign.Saltwatercorrosion andthesubtropicalclimateofourstudyareacausedmultipleelectroniccomponentfailuresduringtheinitialmonthsof setupandoperation.Thesefailureswerere”ectedinthelowand variablecaptureprobabilities( p )immediatelyafterconstruction. Afteradaptingantennaecomponentstoenvironmentalconditions,themajorityofelectronicfailuresceasedand p increased. Anotherdesignchallengewas“ndingsuitableareastoplaceantennaeformaximumresightingef“ciency.Thedimensionsofthe copperinductorcoilandthereadrangeofthemagnetic“eldlimitedantennaapplicationtolocationswhere“shmovementwas restrictedtonarrowareas<10…30minwidth(e.g.,narrowpassages increeks)asopposedtoopenhabitats,suchasbays. Seinenettingalsoprovidedadvantagesandchallengeswhen comparedtoPITtagantennae.Seinenettingmorethoroughlysampledtheentirestudyarea,albeitduring“nitetimewindows,while PITtagantennaeonlysampled“xedlocations.Samplingmultiple spatialpointsisespeciallyimportantifthestudyspeciesexhibits asmallhomerange( KramerandChapman,1999 ),asanindividual withasmallhomerangemaynotpassstationaryantennae.Also,

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160 A.B.Barbouretal./FisheriesResearch121-122(2012)153…160thephysicalrecaptureofmarkedindividualsprovidedinformation ongrowthandpopulationsize,importantmetricsfordetermining habitatqualitythatantennaeresightingscannotmeasure.However,therecaptureinef“ciencyofseinenetsinoursystemmade collectionofgrowthinformationdif“cultwithoutasupplementary method(e.g.,otolithaging). 4.3.Conclusions Sincephysicalcaptureisrequiredtomarkthestudysubject, theuseofPITtagantennaecanbeconsideredaneffectivepartof adual-methodapproachincorporatingbothphysicalcaptureand telemetry.Theuseofthisdual-methodapproachhasthepotential toadvanceecologicalstudiesofcoastal“sh.Althoughtheusefulnessofdatafromtheseinenetswaslimitedbylowrecapturerates andenvironmentalconditions,PITtagantennaefunctionedcontinuouslyandrecapturedmarked“shathighrates.Likewise,PITtag antennaecouldnotcollectgrowthorpopulationsizedata,while seinenetrecapturescould.Therefore,wearguethatwhenusedin conjunction,thetwomethodscanprovideamorecompletepictureof“shhabitatuseandsurvival,therebymakingastronger contributiontounderstandinghabitatusebyestuarine“shes. Althoughnotcomparedinthisstudy,PITtagsalsohavelonger lifespans,arelessexpensive,andareofferedinsmallersizesthan othertelemetrytags,suchasacoustic,GPS,orradiotransmitters. Whiletheseothermethodscanbeusedinawidervarietyofhabitats thanPITtagantennae,thecostofthesetagswouldmakeitdif“cult tomarkahighnumberofindividualsandbatterylifewouldprohibit long-termtrackingofspeci“cindividuals.Thus,thecombineduse ofPITtagantennaeandphysicalcapturemaybeanidealapproach forcost-ef“cient,long-termstudiesofcoastalorestuarineanimals. Acknowledgments WethankM.Allen,M.Lauretta,K.Lorenzen,andW.EPineIIIfor commentsonthemanuscript.WethankDonandDorothyGulnac, R.Boucek,andN.Barkowskiforhelpconstructingantennae.This publicationwassupportedbygrantsfrom:theNationalSeaGrant CollegeProgramoftheU.S.DepartmentofCommercesNational OceanicandAtmosphericAdministration(NOAA),GrantNo.NA06OAR4170014toAJA;CharlotteHarborNationalEstuaryProgramto AJA;SouthwestFloridaWaterManagementDistricttoAJA;bya NationalScienceFoundationGraduateResearchFellowshipunder GrantNo.DGE-0802270toABB;andbyaNSFResearchExperienceforUndergraduateFellowshiptoTY.Theviewsexpressedare thoseoftheauthorsanddonotnecessarilyre”ecttheviewofthese organizations. ReferencesAdams,A.J.,Hill,J.E.,Kurth,B.N.,Barbour,A.B.,2012.Effectsofaseverecoldevent onasubtropical,estuarine-dependent“sh.GulfCarib.Res.24,submittedfor publication. Adams,A.J.,Wolfe,R.K.,2006.Cannibalismofjuvenilesbyadultcommonsnook ( Centropomusundecimalis ).GulfMex.Sci.24,11…13. 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