Postrelease Mortality of Angled Tarpon in Puerto Rico by Puerto Rico Sea Grant

PostreleaseMortalityofAngledTarponinPuertoRico

LauraB.Horowitz,PeterJ.Allen ,J.WesleyNeal,andSandraB.Correa* DepartmentofWildlife,FisheriesandAquaculture,MississippiStateUniversity,Box9690,MississippiState,Mississippi 39762,USA

Abstract

Tarpon Megalopsatlanticus isapopularandeconomicallyimportantinshoresport fishinPuertoRico,andthe pursuitofthisspeciesbylocalanglersandtouristscontributestotheeconomy.Thisspeciesismanagedasano-take fishery,whichaimstopreservepopulationsbycatchingandreleasing fishthatwouldotherwisebesubjectedtoharvest andremovalfromthepopulation.Theapproachassumesminimalmortalityand/orreduced fitnessofreleased fish,yet theprocessofanglingcanproducemanysublethalsideeffectsordirectmortality.Inthisstudy,charteranglingfor TarponintheSanJuanlagoonsysteminPuertoRicowasexaminedtodeterminepostreleasemortality.Angled fish wereexternallytaggedwithacoustictransmittersandrelocatedperiodicallytodeterminethefateafterrelease. Postreleasemortalitywasatleast4.5%(two fishobserveddead)andatmost18.2%(observeddeadplusclassified deadbasedonlackofmovement;confidenceinterval[CI] = 7.5–28.9%).Sometaglosswasobserved,whichcould havearti ficiallyelevatedclassifiedmortalityestimates.Potentialfactorsthatcontributetomortalityarediscussedand comparedtotheliterature.Hooktype,gearaction,landingprocedures,andairexposurewerekeyareasofpossible improvement.Recommendationstominimize fishharmduringanglingincludetheuseofheavieractiongeartoreduce fighttime,acirclehookrequirementforlivebaittoreducedeephooking,maintaining fishinthewaterduringlanding andphotography,andlimitingairexposureto2minorlessif fishareremovedfromthewater.

Thegoalofcatch-and-releaseanglingisthatcaptured fishsurvivetobecaughtagain(PollockandPine 2007). Thisrequiresthat fi sharenotphysiologicallycompromisedafterrelease(Orr 2009).Despitetheintentofcatchand-releasepractices,anglingcancauseacutestressand physicaldamageto fi sh,withpotentiallethalorsublethal consequences(Mazeaudetal. 1977;Skomal 2006).Visible injuriesmayincludehookingwounds,physicaldeformities fromrepetitivehookingandhandling,andbacterial/fungal infections(Meka 2004).Subsequenttocapture, fishcan experiencerespiratoryissuesandcompromisedimmune systems,leadingtobehavioralchanges(Cookeetal. 2002; Wilsonetal. 2014).Otherphysiologicalimpairments includedamagetothere flexresponse,increasedriskof predation,anddecreasedindividual fitness(Campbell

*Correspondingauthor: sbc257@msstate.edu ReceivedMay25,2022;acceptedJanuary14,2023

etal. 2010;Brownscombeetal. 2017).Thesestressors mayactindividuallyorsynergisticallytoresultinthe deathofreleased ﬁsh.Mortalitymayvarybyduration andmagnitudeofthestressors(MekaandMcCormick 2005)andareinﬂ uencedbygeartype(Sassetal. 2018),environmentalconditions(Suskietal. 2006),and postcapturehandlingandreleasepractices(Guindon 2011).

Acoustictelemetryisoftenusedtodetermineinsitu mortalityoveraperiodofhourstoweeksfollowing release(Princeetal. 2002;FFWCC 2013).Forexample, RedSnapper Lutjanuscampechanus taggedwithacoustic transmittersdemonstratedanglingmortalityoccurred within72hfollowingrelease(EbertsandSomers 2017). ResultsfromacoustictelemetryofBone ﬁsh Albulavulpes

Thisisanopenaccessarticleunderthetermsofthe CreativeCommonsAttribution License,whichpermitsuse,distributionand reproductioninanymedium,providedtheoriginalworkisproperlycited.

suggestedangling-inducedmortalityoftenoccurswithin minutesof ﬁshbeingreleased(Danylchuketal. 2007). Furthermore,43%of30Bone ﬁshsuccumbedtoanglinginducedpredationwithina2-weektimespan(Moxham etal. 2019).Thesestudiesutilizedexternalattachmentof transmitterstoavoidintrusivesurgicalprocedures. Althoughexternaltagginghasbeenfoundtoreduce growthandlong-termsurvival,directmortalityfrom externaltaggingisrare(seereviewbyJepsenetal. 2015).

Tarpon Megalopsatlanticus isapopularinshoresport fishspecies,andthispopularityderivesfromitslargesize, powerful fightwhenhooked,andaccessibility,asTarpon aremainly fishedinshoreandincalmcoastalwaters (Guindon 2011).InPuertoRico,thepursuitofthisspeciesbylocalanglersandislandvisitorscontributessignificantlytotheeconomy(Garcia-Molineretal. 2002). However,Tarpon fisheriesinPuertoRicopreviouslywere pronetopopulationdeclinesduetoanglingandharvest (Zerbietal. 2001).Between2000and2003,upto10,189 kgofTarponwereharvestedannually,withnegative effectsonpopulationsize(GuerreroPérezetal. 2013).In response,thePuertoRicoDepartmentofNaturaland EnvironmentalResources(DNER)imposedaharvest prohibitiononTarponin2004toprotectstocksfor anglingbylocalsandtourists(GuerreroPérez etal. 2013).Currently,Tarponanglingisonlycatchand releaseinPuertoRicobylaw.

Dataoncatch,anglingeffort,andpostreleasemortalityofTarponhavenotbeencollectedinPuertoRico (GuerreroPérezetal. 2013).Guindon(2011)and Edwards(1998)reportedlowpostreleasemortalityin SouthFlorida,buttheresultsmaydifferfor fishangled inotherlocations.Tarponarewidelydistributed(WinemillerandDailey 2002),andenvironmentalparameters (i.e.,waterqualityandchemistry;Leichteretal. 2006) andvaryinganglingtechniques(Brownscombe etal. 2017)mayinfluencethepostreleasefateof fish. Therefore,thisstudyevaluatedcatch-and-releasemortalityofrecreationallyangledTarponinPuertoRicousing acoustictelemetry.Thespeci ficresearchobjectivewasto evaluatethefate(aliveordead)ofTarponusingacoustictelemetry.Factorsinfl uencingpostreleasefatearediscussed,andthisinformationcanbeusedtodecreasethe impactofcatch-and-releaseanglingontheTarpon fishery.

METHODS

Studysite. SanJosé,LosCorozos,LaTorrecilla,and PiñonesLagoonandassociatedinterconnectedcanalslie withinthemetropolitanareaofSanJuan,PuertoRico (Figure 1)andarecollectivelyidentiﬁ edasoneofthe world'stopTarponanglingdestinationsbythemedia. Tidalcyclesinﬂuencewaterqualityandsalinity,asdoes

rainfallinputviafeederstreamsandcanals.Lagoonsare surroundedbyredmangrove Rhizophoramangle,black mangrove Avicenniagerminans,andwhitemangrove Lagunculariaracemosaforests(Pooletal. 1977).Dueto dredging,landfills,andotheranthropogenicactivities,the hydrologicalcharacteristicsoftheestuaryhavechanged considerablyoverthepastcentury(Villanueva etal. 2000).Hydrologicalcharacteristicsthathavebeen alteredbyurbanizationintheSanJuanlagoonsystem includedisconnectionfromSanJuanBay,sedimentdenitrification,and fluxesofammonium,nitrate,andphosphorusnutrients(Pérez-Villalonaetal. 2015).These modi ficationshaveresultedinarelativelyclosedsystem withasinglesmallconnectiontotheAtlanticOcean throughthenarrowpassageofBocadeCangrejos,inthe localityofPiñones.

Anglingandtelemetry. Thisresearchwasconducted inpartnershipwithalocalchartercompanythatspecializesinSanJuanlagoonTarponangling.Theguides hostedresearchersoncharteredtripswithclientsfrom MaytoJuly2021.WhenaclienthookedaTarpon,informationabouttheanglingprocesswasrecordedfrom hooksetuntillanding.Here,theanglingprocessincluded fightduration,thenumberofjumps,rodaction(light, medium,orheavy),andhandlingondeck(i.e.,hook removalandbodysupportforphotography).Wealso recordedthegeographiclocationofcapture.Tarponwere onlycapturedusingspinningorbait-castinggear,withlive baitoncirclehooksduringthestudy.Tolandeach fish, theguidewouldgrabthe fishbythelowerjaw(hand lipped),and fishwerepulledintotheboatoverthegunwaleorbowandontothedeck.Dataonlandingconditionandproceduresincludedhooklocationonthe fish, observedinjurysuchasbleeding,andhandlingprocedure duringphotography.Onceclientsweredonewiththeir captured fi sh,itwashandedovertotheresearchersfor measurementoftotallength(mm)andpossibletag implantationpriortorelease.WerandomlyselectedTarponfortagimplantation.For fishnotselected,bloodwas drawnforanotherstudypriortorelease.

Transmitterimplantationandtelemetry. Acoustictag assembliesconsistedofaSonotronics(Tucson,Arizona) (CT-82-2-E)transmitterattachedtoatitaniumanchor(Ti Small,WildlifeComputers,Redmond,Washington)with astainless-steelleader(42.2kgtest,0.005cmdiameter) andcrimpsleeves(doublebarrelsize7,0.135cmdiameter).Thelengthoftheleaderwasadjustedto fishsizeso thatthetransmitterdidnotreachtoorinterferewiththe dorsal fi n.Tagassemblieswereattachedtothe fish'sleft side,twoscalerowsbelowtheinsertofthedorsal fin. Usingforceps,twotofourscaleswereremovedfromthe locationoftaginsertiontofacilitateimplantationofthe anchor.Thetaganchorwasinsertedwithatagpole(comprisedbyAZ-TAGPOLE-004;SwobbitTaggingPole

FIGURE1.Mapof (A) thelocationofPuertoRicointheCaribbeanSea, (B) thelocationoftheSanJuanlagoonsysteminPuertoRico,and (C) a close-upoftheSanJuanlagoonsystemandsurroundingareawiththelocationoftrackingsitesandpassivereceivers;BocadeCangrejosconnectsthe systemtotheAtlanticOcean.MapswereproducedbyKaroldCoronado-Franco.

16.51cmlength,AZ-DARTBUSH-001;DartBushing ﬁ ts 1.91cmpole,andAZ-DARTAPP-011;DartApplicator1/ 4–28;WildlifeComputers)withinthescaleremovalzone throughthemusculatureandbetweenthepterygiophores. Fishwereimmediatelyreleasedbacktothesiteofcapture.

Relocationeffortswereconductedusingactivetracking onceperweekwithaSonotronicsManualTrackingKit (MANTRAK;whichcontainsaUSR-14receiver,DH-4 directionalhydrophone,TH-2towedomnidirectional hydrophone,andaccessories).Priortothestudy,researchersestablished28listeningstations,eachselectedtooptimizelagooncoveragebyutilizingamaximumlineofsight distanceofnomorethan1kmbetweenlisteningzones (Figure 1).Duringeachtelemetryperiod,theboatwas

maneuveredtoeachlisteningstationandtheomnidirectionalhydrophonewasloweredintothewaterandallfrequencieswerescanned.Whenatagged ﬁshwasdetected, approximatelocationwasdeterminedusingthedirectional hydrophone.Whenalltagfrequencieshadbeenscanned, theboatwasmaneuveredtothenextlisteningstation.

BecauseTarponarehighlymobile(e.g.,Duffi ng Romeroetal. 2021),thelagoonsystemislargeandcomplex,andtimetoconducttelemetrywaslimitedto1dper week,exactlocationsusingtriangulationwerenotdetermined.Instead,theprotractedtelemetryperiodwasused tocon firmwhethera fishwasnonmobileandpresumed deadormobileandalive.Inthisstudy,postreleasefate wasassessedbyrelocating fishweeklyforupto9weeks. Thepostreleaseevaluationperiodwassetat5d,soto

classifya ﬁshasalive,movementmustbedetected5dor moreafterrelease.

Classifi cationrulesweredevelopedtointerpret fishstatus basedonmovement(seeFigure 2 forexamples).Ruleswere basedonpreviousstudiesthatinferred fi shstatusbyclassifyingandgroupingpatternsoftelemetrydatafromreceivers (HeupelandSimpfendorfer 2002;Villegas-Ríosetal. 2020; Weinzetal. 2020).Fora fishtobeconsidered “alive,” it mustmeet allthree ofthefollowingcriteria(Figure 2B):

1. ﬁshwererelocatedattwoormoredifferentlistening stationsatleast1weekapart;

2.observedmovementoccurredatleast5dafterrelease (indicatingitwasalivethrough5d);and

3.listeningstationscannothaveoverlapping ﬁ eldsof detection(i.e., >1km).

Fora ﬁshtobeconsidered “dead,” itmustmeet any of thefollowingcriteria(Figure 2C):

1.duringallweeklydetectionsafterday5withaminimumofthreedetections, ﬁshwasdetectedinthesame area(withinthe1kmreceiverrange);

2.researchersrecoveredthedead ﬁsh;or

3.researchersreceivedreportsofa ﬁ shbeingdeadwithin the5-devaluationperiod,anditwasrelocatedinthe reportedlocationfortwoormoreweeklytracking periods.

FIGURE2.Examplesof fishmovementandstatusclassificationasalive,dead,orcensoredbasedonrules. (A) Comparativemapofallthree classificationsrepresentedbypanels(B–D). (B) Fishconsideredalivebecauseofsignificantlymovementafter5d. (C) Fishconsidereddead,asit movedfromtheinitialsiteofcapturebutdidnotmovesignificantlyfollowingthe5-devaluationperiod. (D) Fishunabletobeclassifiedandcensored fromanalyses.MapswereproducedbyKaroldCoronado-Franco.

Fora ﬁ sh'sstatustobeunknownandthedata “censoredfromanalyses, ” itmustmeet any ofthefollowing criteria(Figure 2D):

1.the ﬁshwasfoundjustonce(indicatingitcouldhave leftthesystem,experiencedtagfailure,etc.);

2. ﬁshhadonlytwodetections,withthe ﬁ rstdetection occurringwithinthe5-devaluationperiod;or

3. ﬁshhadonlytwodetectionsinthesamelocationor overlapping ﬁeldsofdetectionimmediatelyfollowing the5-devaluationperiodonconsecutivetrackingdays andthendisappeared.

TwopassiveSubmersibleUltrasonicReceivers(SURs; Sonotronics)wereplacedatthemouthofthelagoonsystem todetermineif ﬁshexitedtotheAtlanticOcean(Figure 1). Beforedeployment,passivereceiverbodies(excludingthe transducer)werewrappedinblackplasticandelectrical tapetoreducebiofouling.MountswereconstructedofPVC poles(5.08cmdiameter × 1.22mheight)insertedinconcretebases;PVCwasspray-paintedblackforconcealment. ReceiverswereattachedtoPVCmountswith550paracord andnyloncableties.Oncedeployed,SURmountswere tetheredtodockpylonstopreventlateralmovement.

Dataanalysis. SurSoftDPCsav1.0.8(c1)software (Sonotronics)wasusedtodownloadandanalyzepassive receiverdata.Detectionsfromthesereceiverswerecombinedwithactivetelemetrydatafortheassessmentof fish status.Combined fishlocationdatawereplottedinArcMap(10.8.1)toanalyze fi shmovementfordetermination of fi shstatus.Satellitebasemapswereretrievedfromthe ArcGISwebsite(ESRI; https://www.esri.com/).Confi dence intervalsformortalitywerebasedonmethodsusedby Wilde(2002)todeterminecon fidenceofstatusaround fish usingacoustictracking.

Summarystatisticsincludedall fishcaughtduringthestudy (includingthosenotreceivingtransmitters).Linearregression assessedwhether fishsizeinfluencedthedurationofthe fight. Priortoregression,theresponseandpredictorvariableswere log-transformedtocorrectfornormalityandtolinearizethe relationship.RegressionwasimplementedinR(version4.1.3; RFoundationforStatisticalComputing,Vienna).

RESULTS

AnglingProcess

Anglingprocedureswereevaluatedfor93angledTarponthatrangedfrom47to200(mean = 88)cmintotal length.Fishsizedirectlyinﬂuenced ﬁghttime(adjusted r 2 = 0.55, P < 0.0001)(Figure 3).Fighttimerangedfrom 1to41minbutwasskewedtowardsshortertimeswitha mean(±SD)of5(±7)min.Observedjumpsrangedfrom 0to14(mean ± SD = 3 ± 2).

FIGURE3.Linearregressionshowingtherelationshipof ﬁshtotal length(cm)with ﬁghttime(min).

Thechartercompanywithwhomwepartneredprimarilyusedlightactionrods,livebait,andcirclehooksfor charteredanglingtrips(Figure 4).Withcirclehooks,most fishwerecaughtineitherthecornerofthemouthor upperjaw,andmost(90%)werereleasedafterthehooks wereremoved;however,whenhookingoccurredinthe throatordeeper,lineswerecutleavingthehookinplace toreduceinjury.Littletonobleedingwasobserved amongmostangled fi sh(Figure 4).Followinghook removal,clientscouldtakephotographswiththe fi shand choosethepositiontoholdthe fish.Horizontalphotographywasthemostpopularandencouragedbytheguides tobettergraspthe fish(Figure 4).Thispracticehadthe clientsupportthe fish'sbodyweightwithonehandonthe abdomenwhilecontrollingthe fishbyholdingthelipwith theotherhand.Larger fishwereoftenphotographedlying onthedeck.

PostreleaseMortality

TaggedTarponmovedconsiderablybetweenlagoons, suggestingthattheclassi ficationrulesbasedonmovement arevalid.Three fishweredetectedatthelagoonentrance, butno fishexitedtotheAtlanticOceanduringthestudy. Ofthe49tagged fi sh,5werecensoredfromfurtheranalysesduetoinsuffi cientdatatoclassifya fishasaliveor dead.Oftheremaining44classifi ed fi sh,36wereclassi fied asalivebasedonobservedmovementfollowingthe5-d evaluationperiod(81.8%).Conversely,eight fishwere classifiedasdeadbasedonlackofmovement(18.2%).

FIGURE4.Factorsduringtheanglingprocessthatcouldinﬂuencemortality.Rodaction(topleft),hookinglocation(topright),observedbleeding (bottomleft),andhandlingpositionduringphotography(bottomright)arepresented.

One fishdiedsoonafteranglingandthetransmitterwas retrieved;another fi shwasreported fl oatingdead,and althoughnotphysicallyrecovered,itwasrelocatedconsistentlyintheareareported.Thisyieldsacon firmedmortalityof4.5%(fishencountereddead),andapotential mortalityofupto18.2%estimatedusing fishstatuscriteria(fishconsidereddeadbasedonpostreleasemovement, includingcon firmedmortality).TheCIofthemortality estimateusing fishstatuscriteriawas7.5–28.9%.

DISCUSSION

Thisstudywasthe firsttoinvestigatecatch-and-release mortalitywithintheTarpon'stropicalrange.Mortality within5dofanglingwasatleast4.5%andatmost 18.2%,basedoncon firmedandclassi fiedmortalities, respectively.Thetwocon firmedmortalitiesoccurred within24hofrelease.Thissuggeststhatpostreleasemortalityoccursquickly,andthe5-devaluationperiodwas adequatetoassesstheeffectofangling.Itispossiblethat someclassifiedmortalitiesrepresentedtagshedding insteadof fi shdeathduringthe5-devaluationperiod,as thelossrateofexternaltransmitterscanbesubstantial

andhasbeenreportedtobeashighas100%insome studies(seereviewbyJepsenetal. 2015).TwoTarpon werereportedlycapturedbyotherchartersthathadscarringconsistentwithtagloss,andathirdwascaptured withthetetherinplacebutthetransmittermissing,lendingcredencetothisassertion.Thisstudydidnotcollect dataontagretention.Taglossthatoccurredaftertheinitialclassiﬁ cationperiod(afterpostreleasemovementwas conﬁ rmed)wouldnotaffecttheresults.

Therangeofpostreleasemortalityfromthisstudy wascomparabletooneestimatefromFlorida,where Guindon(2011)reporteda13%mortalityrate(95%CI: 6–21%)inangledTarpon(n = 82).Thatstudydescribed predationonreleasedTarponbysharkspeciestobea majorvectorofmortality(8.3%).Sharkpredationratesof nearly28%wererecordedinanotherlarge-scalesatellite taggingstudyofTarpon(n = 292)extendingfromNorth CarolinathroughouttheGulfofMexico(Luo etal. 2020).Suchahighpredationratebysharksmay, however,beanartifactofthestudysincesharksarelikely attractedtotheelectric ﬁeldgeneratedbysatellitetags andthepredationrateincreasedinareaswithhighshark density(Luoetal. 2020).Thecurrentstudydidnot

observepredationonTarpon,andpredatorysharkspecies arenotcommoninthelagooncomplex.Anotherstudy fromFlorida(n = 27)reportedlowermortality(3.7%)of angledTarpon,andtheauthorsattributedthislowrateto anglingpractices,includingstyleandsizeofhook,useof heavygear,andaggressiveanglingtechniquestobringthe ﬁshtotheboatasquicklyaspossible,andreleasingthe Tarponbynottakingitoutofthewater(Edwards 1998). Anglinginthecurrentstudyutilizedcirclehooks,which reduceddeephooking.However,lightactionspinning rodsweretheprimarygear,whichprolonged ﬁghttimes andlikelyincreasedthetimeofphysicalandphysiological stress(Rothetal. 2018).Likewise,thePuertoRicocharter guidestypicallyremoveTarponfromthewaterforlandingandphotography,inducingadditionalstressandair exposure.

Certainfactorshavebeenidenti ﬁedintheliterature thatreducemortalityduringtheanglingprocess.Hooking locationcanaffectanglingmortalityinTarpon(Guindon 2011)andotherspecies(e.g.,RainbowTrout Oncorhynchusmykiss,Meka 2004).Potentiallylethalhooking locationscanincludetheeye,gills,andesophagus (Ostrandetal. 2005).Forexample,Derbio Trachynotus ovatus hookedinternallyhadan85%mortalityrate(Alós etal. 2008).Similarly,95%and75%ofSpottedSeatrout Cynoscionnebulosus diedwhenhookedintheesophagus andgills,respectively,comparedtoonly10%when hookedinthebonymouth(Jamesetal. 2007).Theuseof circlehooksinthecurrentstudycontributedtotheprevalenceofnonlethalhookinglocations(Cookeetal. 2003).

Most(n = 78)Tarponwerehookedinthecornerofthe mouthorintheupperorlowerjaw,withfew(n = 15) ﬁsh hookedinotherlocations(i.e.,outerheadorbody,inner oralcavity,oresophagus).

Most(91%)ofthe fishhadthehookremovedpriorto release.Onlyhooksthatweretoodeeptosafelyremove wereleftinthe fishbycuttingtheline.When fishhave deeplyingestedhooks,itisoftenmorebene ficialtoleave thehookintheindividual,asattemptingtoremovethe hookoftenresultsinadditionalinjuriesandincreased mortality(Butcheretal. 2007).InWhiteSeabass Atractoscionnobilis,forexample,leavingdeeplyembedded hooksincreasedsurvivalrates,and39%ofhooksthat weredeeplyingestedcouldpassthroughtheorganism's system(Aalbersetal. 2004).Whenlookingatsimulated anglinginSandWhiting Sillagociliate,23%of fishthat ingestedahookdied,butsurviving fishwereabletofeed, andsome fi shpassedingestedhookswithlimitedlongtermphysiologicalimpactsdocumented(McGrath etal. 2009).

Hookinglocationcandeterminewhetherbleeding occurs,especiallywhenthehookpenetratessoftorhighly vascularizedtissuessuchasthegillsandesophagus. Althoughsomebleedingwasobservedinthisstudy,76%

oftheindividualsexperiencednobleeding,withonlytwo individuals(2%)experiencingheavybleeding(gillhooked), oneofwhichwascon ﬁrmeddead.Previousstudieshave notedbleedingintensitywithhookinglocationcausing increasedmortality(e.g.,SchislerandBergersen 1996). Conversely,hookinglocationdidcorrelatewithbleeding intensitybutnotmortalityinArcticGrayling Thymallus arcticus (Casselman 2005).

Rothetal.(2018)assertedthatlonger fighttimesmay causephysiologicaldisturbancesthatcanleadtoagreater riskofpostreleasemortality.Increased fighttimeinShortfinMako Isurusoxyrinchus didincreasephysiological stressfactorssuchaslactateandglucosebutdidnot impactsurvival(Frenchetal. 2015).ForBrookTrout Salvelinusfontinalis,mortalitywasindependentof fight time(Kerretal. 2017).Similarly,Guindon(2011) reportedtheaverage fi ghttimeofTarponwas23.7min for fishthatsurvivedand16.5minfor fishthatexperiencedmortality,whichiscontrarytoRothetal.'s(2018) hypothesis.Themean fi ghttimeinthecurrentstudy(5 ± 7min)was3–5timeslessthantheGuindon(2011)study, despiteanglingprimarilyusinglightactiongear.Rod actionrelatestohoweasilyandwherealongtheshafta rodbendswhentensionisappliedtothetip.Manystudies previouslyhavenotedtheactionofthegearusedbutdid notrelateittoanglingmortality(Jones 2005;Danylchuk etal. 2014;Holderetal. 2020).Catchingthe fi shfaster reducestheintensityofthe fi ghtandstresstothe fi sh,but chartercompaniesmaypreferalongeranglingexperience forclients.Loweringtheactioncreatesmoreexcitingand challenging fi ghtforcustomers.

Thedisparityin fi ghttimebetweenGuindon(2011) andthecurrentstudywaslargelydueto fishsize. AlthoughlargerTarponarecapturedintheSanJuan lagoonsystem,the fisheryislargelysupportedbysmaller fish.Tarponevaluatedduringthecurrentstudyaveraged 88cmtotallength(±27cm),whereas fishintheFlorida evaluationaveraged160.8cmfor fishthatsurvivedand 146.5cmfor fi shthatexperiencedmortality(Guindon 2011).Thelackofarelationshipbetween fishsize andpostreleasemortalityreportedbyGuindon(2011)was furthersupportedbyameta-analysisofthecatch-andreleaseliterature(BartholomewandBohnsack 2005).This isasurprising findingbecauselarger fishtendtohave greater fighttimesandpresumablymoreangling-induced stressresponse(Guindon 2011).Fishsizewaspositively correlatedwith fi ghttimeandhandlingtimeinthecurrent study,andotherstudiessupportthis finding(e.g.,Danylchuketal. 2014;Pinderetal. 2017).ForBrookTrout, fighttimewasnotcorrelatedwithtotallength(Kerr etal. 2017),suggestingtherelationshipof fighttimeand lengthismostlikelyspecies-dependent.

Tarponcanbeacrobaticandleapfromthewater duringangling,attemptingtodislodgethehook

(Kokomoor 2010).Inacon finedmangroveenvironment liketheSanJuanlagoonsystem,jumpingbehaviorhas potentialtoinjureviaimpactwithmangrovesandother obstacles,orwiththewatersurface,butthishasnot beenempiricallyestablished(LuoandAult 2012).Inthis study,Tarponaveraged3(±2)jumpsduringtheangling process.Schlenkeretal.(2016)observedthatjumpingby WhiteMarlin Kajikiaalbida increases fishexhaustion andaddsadditionalstresstotheindividual(Schlenker etal. 2016).Further,jumpingbyangledRainbowTrout correlatedwithincreasedriskofinjury,asjumpingoften ledtodeeperhookwoundsandentanglementofthe fish intheline(Meka 2004).Itislikelythatjumpingbehaviorattheveryleastaddstothestressresponseduring angling.

Landingisarelativelystressfulprocess,particularly forlargerTarpon,whichweredraggedbythelowerjaw overtheedgeofthegunwaleorbowandintothe flooror deckofthevessel.IntheevaluationbyEdwards(1998), Tarponweresupportedinthewater,thuseliminatingair exposureandrelatedinjury,andsurvivalrateswerehigh, at96.3%.AlthoughTarponarefacultativeairbreathers (Geigeretal. 2000),thisraisesthequestionofwhether Tarponhavegreatersurvivalwhenleftinthewaterto preventairexposureandtheeffectsofgravityoutsidea liquidmedium.Watersupportstheweightofa fi sh,and removalfromthewatercancauseinjurytointernal organs,mandibularbonebreaksandseparatedtongue (Danylchuketal. 2008),andvertebralseparation(Gould andGrace 2009;Frawley 2015),andunexpected fish movementcanleadtoaccidentaldropsandimpactinjuries.Increasedairexposurehasbeencorrelatedwith increasedstressin fish(FergusonandTufts 1992;Brownscombeetal. 2017)andanincreasedamountoftime requiredforthe fishtorecoverfromthecaptureevent (Cookeetal. 2001;Brownscombeetal. 2017).Guindon(2011)suggestedlimitingairexposureofTarponto2 minorlessandreportednosigni fi cantphysiologicaldifferenceinbloodchemistryresponsewith1minairexposure.Whereasevidencesuggestslongerairexposurehas adverseeffectsonthesurvivabilityofsomespecies (ArlinghausandHallermann 2007),itmightbeadvantageoustominimizeairexposure,assuggestedby Edwards(1998)forTarpon.

Anglersliketosharetheirexperiencewithothers,especiallythroughphotography.Clientswereencouragedto hold fi shhorizontallysupported(58%)whentakingphotographs;onlyafew fish(13%)weresupportedvertically oronthedeck,particularlyforlarger fishduetotheir heavyweight.All fishweregentlyrevivedandreleased backintothewateraftercapture.Horizontallysupporting andgentlyreleasingthe fi shminimizesinjuryandstress (Frawley 2015).Practicingtheseeffortsarenowconsideredetiquetteandpartofsafehandlingpracticesofcatch-

and-releaseangling.Someresearchersrecommendleaving the ﬁ shinthewaterforhookremovalandphotography, minimizingairexposureandpossibleinjury(Cookeand Sneddon 2007).

ItiscommonforTarpontoshow “partialmigration,” thatis,intraspeci ficvariationinmigrationdistance,andthespeciescommonlyresidesinestuarine habitatforpartoftheyear(Luoetal. 2020).The uniquenessoftheSanJuanlagoonnetworkisthat Tarponarecontainedwithinasemiclosedsystemand canonlyenterandexitviaashallowopening(Bocade Cangrejos,Figure 1C)thatisabout35mwide.Itis unclearwhetherTarponintheSanJuanlagoonnetwork areyear-roundresidentsormigrateoutofthesystem duringsomepartoftheyear.Inthisstudy,onlythree fishapproachedtheexittotheAtlanticOceanbutdid notleaveforanysignifi cantdurationwithinthe9-week observationperiod.If fishareyear-roundresidents,it raisesconcernabouthowcontinuousanglingpressure impactsindividualsconfi nedwithinasemiclosedsystem. Althoughtheanglingeffortwasnotmeasured,itissubstantial,withmanycharterout fitswithclientanglers andprivaterecreationalanglersusingtheirpersonal watercraftdaily.Itwascommonduringthisstudytobe targetingTarponwithinasmallareawith fi veormore otherboatsnearby.Further,telemetered fishfromthis studywerecaughtmultipletimesbydifferentangling charters,asresearchersreceivedmultiplereportsof tagged fishbeingcaught.Whilethisindicatesthatsome fishrecoverquicklyandreturntonormalfeedingbehavior(Cookeetal. 2013),italsosuggeststhat fi shinthis lagoon fisheryarerepeatedlyexposedtotheriskof injury,airexposure,andpostreleasemortality.

Cautioniswarrantedwhenusingtheuppermortality rateestimates,duetotheclassifi cationprocess.Immobile fishwereclassi fiedasmortalitieswithoutveri fication. Thereweretworeportsof fi shcapturedthatlookedlike thetagwaspulledoutandanotherthatwasdisconnected fromtheanchorline.Thisindicatesthattagshedding occurred,butthetimingandeffectiscurrentlyunknown. Taglosssoonaftertaggingwouldleadtopotentialfalse classificationsifthose fishsurvivedaftersheddingthetag. Further,thisresearchoccurredonlyduringthesummer. Althoughenvironmentalconditionsarerelativelystablein PuertoRico,someseasonalvariationoccursandcould influenceseasonalresults.Summerwatertemperaturesare greaterandthereforemayrepresentaperiodofgreater thermalstress.Finally,thisstudyassessedspeci fi cpracticesofasingleanglingcharteroperator.Theanglingand handlingtechniquesofotherchartercompanies,freelancingcaptains,andindependentanglersmaydiffer(Brownscombeetal. 2017).

Withoptimalanglingandhandlingpractices,catchand-releaseanglingcanbesuccessful,withminimal

mortality(Brownscombeetal. 2017).Anglertechniques canbealteredtominimizecatch-and-releaseangling fromhavingnegativeimpacts(BartholomewandBohnsack 2005).Hooktype,gearaction,landingprocedures, andairexposurewerediscussedasareaswheremodi ficationscouldyieldasigni ficantimprovementinpostrelease survival.Theuseofheavieractionrodsandgreaterline testcouldreduce fi ghttime,shorteningtheperiodof anglingstress(Mohanetal. 2020).Thismaynotbethe bestoptiontomaximizetheanglingexperience,asanglers pursueTarponforthe fight.Circlehooksarewellresearchedtoreducedeephooking,andacirclehook requirementwhenusingnaturalbaitcouldincreasepostreleasesurvival(seereviewsbyCookeandSuski 2004; Kelleretal. 2020).Maintaining fishinthewaterduring landingandphotographywouldeliminateairexposure andinjuriesrelatedtolifting fishfromthesupportwater provides(Edwards 1998).Ifthe fi shmustberemoved fromthewater,itisnecessarytoprovidesupporttothe bodyandreducepotentialforinjury.Finally,whenTarponareexposedtoair,theexposuretimeshouldbekept at2minorless(Guindon 2011).Althoughmostreleased Tarponsurvived,anglingpressureintheSanJuanlagoon systemisintenseandtheserecommendationscouldhelp reducetheriskto fishthataresubjectedtomultiplecaptureevents.

ACKNOWLEDGMENTS

Theauthorsthankourpartnerchartercompany,CaribbeanFishingAdventures,includingA.Muntanerand guidesL.Valdivieso,Y.Sierra,G.Rodriguez,M.Muntaner,andF.Prieto,forworkingwithusthroughoutthe extentofthestudy.K.Coronado-FrancoaidedinGIS, spatialdataanalysis,andmappreparation.Fundingwas providedbyPuertoRicoSeaGrant(Grant2020-2021-007 toJ.W.Neal,P.J.Allen,andS.B.Correa),andadditionalsupportwasprovidedbythePuertoRicoDepartmentofNaturalandEnvironmentalResources,the DepartmentofWildlife,FisheriesandAquacultureat MississippiStateUniversity(MSU),MSUExtensionService,MSUForestandWildlifeResearchCenter,andthe U.S.DepartmentofAgriculture(NationalInstituteof FoodandAgriculture,Grant#1005154toP.J.Allenand Project#MISZ-081700toS.B.Correa).Thisresearch wasconductedincompliancewithprovisionsestablished inMississippiStateUniversityInstitutionalAnimalCare andUseCommittee(Protocol#20-277).Thereisnoconﬂictofinterestdeclaredinthisarticle.

ORCID

PeterJ.Allen https://orcid.org/0000-0003-0122-9387

SandraB.Correa https://orcid.org/0000-0003-4466-6923

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