Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System...
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Evolutionary Applications 20181ndash25 emsp|emsp1wileyonlinelibrarycomjournaleva
Received14June2018emsp |emsp Revised1August2018emsp |emsp Accepted7August2018DOI101111eva12711
O R I G I N A L A R T I C L E
Comparison of coded‐wire tagging with parentage‐based tagging and genetic stock identification in a large‐scale coho salmon fisheries application in British Columbia Canada
Terry D Beacham1 emsp|emspColin Wallace1emsp|emspKim Jonsen1emsp|emspBrenda McIntosh1emsp|emsp John R Candy1emsp|emspDavid Willis2emsp|emspCheryl Lynch2emsp|emspJean‐Seacutebastien Moore3emsp|emsp Louis Bernatchez3emsp|emspRuth E Withler1
ThisisanopenaccessarticleunderthetermsoftheCreativeCommonsAttributionLicensewhichpermitsusedistributionandreproductioninanymediumprovidedtheoriginalworkisproperlycitedcopy2018TheAuthorsEvolutionary ApplicationspublishedbyJohnWileyampSonsLtd
1FisheriesandOceansCanadaPacificBiologicalStationNanaimoBCCanada2FisheriesandOceansCanadaRegionalHeadquartersVancouverBCCanada3DeacutepartementdeBiologieInstitutdeBiologieInteacutegrativeetdesSystegravemes(IBIS)UniversiteacuteLavalQueacutebecQCCanada
CorrespondenceTerryDBeachamFisheriesandOceansCanadaPacificBiologicalStationNanaimoBCCanadaEmailterrybeachamdfo-mpogcca
Funding informationFisheriesandOceansCanadaGenomeBritishColumbiaPacificSalmonFoundationGrantAwardNumberEPIC4GenomeCanada
AbstractWildPacificsalmon includingCohosalmonOnchorynchus kisutchhavebeensup-plemented with hatchery propagation for over 50years in support of increasedocean harvest and conservation of threatened populations In Canada theWildSalmonPolicyforPacificsalmonwasestablishedwiththegoalofmaintainingandrestoring healthy and diverse Pacific salmon populationsmaking conservation ofwildsalmonandtheirhabitatsthehighestpriorityforresourcemanagementdeci-sion-making A new approach to the assessment and management of wild cohosalmonandtheassociatedhatcheryproductionandfisherymanagementisneededImplementationofparentage-basedtagging(PBT)mayovercomeproblemsassoci-atedwithcoded-wiretag-based(CWT)assessmentandmanagementofcohosalmonfisheriesprovidingat aminimum informationequivalent to thatderived from theCWTprogramPBTandgeneticstockidentification(GSI)wereusedtoidentifycohosalmonsampledinfisheries(8006individuals)andescapements(1692individuals)inBritishColumbiatospecificconservationunits(CU)populationsandbroodyearsIndividualsweregenotypedat304singlenucleotidepolymorphisms(SNPs)viadirectsequencingofampliconsVeryhighaccuracyofassignmenttopopulation(100)viaPBT for 543 jack (age 2) assigned to correct age and collection location and265coded-wiretag(CWTage3)cohosalmonassignedtocorrectageandreleaseloca-tionwasobservedwitha40774mdashindividual267mdashpopulationbaselineavailableforassignmentCohosalmonfromun-CWTedenhancedpopulationscontributed65ofthecatchinsouthernrecreationalfisheriesin2017ApplicationofaPBT-GSIsystemofidentificationtoindividualsin2017fisheriesandescapementsprovidedhigh-res-olutionestimatesofstockcompositioncatchandexploitationratebyCUorpopula-tion providing an alternate and more effective method in the assessment andmanagementofCanadian-origincohosalmonrelativetoCWTsandanopportunityfor a genetic-based system to replace the current CWT system for coho salmonassessment
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1emsp |emspINTRODUC TION
WildPacific salmon includingCoho salmonOnchorynchus kisutchhavebeensupplementedwithhatcherypropagationforoverhalfacenturyinsupportofincreasedoceanharvestandconservationofthreatenedpopulationsThepotentialnegativeeffectsofhatcheryproduction and the associated exploitation pressure that may beappliedtonaturalpopulationshavelongbeenrecognizedandhavegainedprominenceinrecentyears(ArakiBerejikianFordampBlouin2008Hilborn1992JonesCornwellBottomSteinampAnlauf-Dunn2018McClureetal2008)leadingtocallsforincreasedresponsi-bilityinmanagementofhatcheryproductionandofthemixed-stockfisheriesitsupports(Flagg2015HSRG2014)InCanadatheWildSalmon Policy (WSP) for Pacific salmonwas establishedwith thegoalofmaintainingandrestoringhealthyanddiversePacificsalmonpopulations making conservation of wild salmon and their habi-tatsthehighestpriorityforresourcemanagementdecision-making(FisheriesandOceansCanada2005)Fisheriesandhatcherysupple-mentation (termedenhancement inCanada)are tobemanaged insuchawayastoensurethatwildpopulationsaresafeguardedandharvestbenefitsaresustainableWildsalmonpopulationsareiden-tifiedandmaintained inconservationunits (CUs) that reflect theirgeographicecologicalandgeneticdiversity
CohosalmonarecaughtincommercialrecreationalandFirstNations fisheries in British Columbia and determination of theimpact of these fisheries is of fundamental importance to statusassessmentforwildpopulationsofconservationconcernandman-agementoflarge-scalehatcheryproductionCurrentandhistoricalassessmentoffisheriesimpactshasbeenconductedwiththeap-plicationofcoded-wiretags(CWTsJeffertsBergmannampFiscus1963)CWTsareappliedtojuvenilefishpriortotheirhatcheryre-leaseandrecoveredfromadultfishheadscollectedfromfisherieshatchery broodstocks and in-river escapement sampling OncerecoveredthetagsaredecodedtodeterminethehatcheryoriginandageoftheindividualfishOriginallyonlycohosalmonmarkedwithaCWTalsoreceivedanadiposefinclippriortohatcheryre-leasewith theexternally visible clipmarkallowingCWT-markedfish to be identified visually and sampled from fisheries or rivercollections
Sincethelate1990sallcohosalmonreleasedfrommanyhatch-eries in southern British Columbia (BC)Washington andOregonhave received an adipose fin clip (termedmassmarking) in orderto facilitatemark-selective fisheries intended to harvest hatcherysalmononlywithmost clipped individuals carrying noCWTThisapproachhasresultedinreducedexploitationofnaturallyspawnedcohosalmonespeciallyinsportfisheriesbutthepresenceofmanyadipose-clippedsalmonwithoutaCWThasimpairedtheefficiency
ofCWT recovery In spite of implementation of an electronic tagdetectionsystemtopre-screenaportionofthecommercialcatchtoidentifysalmonwithaCWTtheprocessingofmanyheadswithouta CWT from voluntary recreational recoveries and the increasingcostsofCWTapplicationandrecoveryhavecauseddegradationofthe informationobtained from the currentCanadian coho salmonassessmentprogram
For coho salmon fisheries impacts are evaluated through theFishery Regulation AssessmentModel (FRAM PFMC 2008) TheFRAMisadiscrete-time-stepage-structuredcomputermodelusedtopredicttheimpactsofavarietyofproposedfisheryregulationsforasinglemanagementyearinimplementationofprovisionsofthePacificSalmonTreaty (PST) related tocohosalmonCanadianandAmericanassessmentstaffusethebilaterallyagreeduponFRAMtoestimatefisheryimpactsPriortoeachfishingseasonbothcountriesincorporatestock-specificconservationconstraintsidentifiedinthecohosalmonchapterofthePST intoFRAMevaluationofplannedfisheriesApplicationoftheFRAMrequiresuseofdataderivedfromfisheryandescapementrecoveriesofCWTsandterminalarearunsizeestimatesforreturnyearsfromabaseperiod(1986ndash1992)Thecurrentexpectedcatchofaspecificstockisestimatedastheprod-uctof(theexpectedabundanceofthestockinafishery)(thebaseperiodexploitation rate)(a correction factor) that relates theex-pectedcatchoreffortinaparticularyearrelativetothatobservedin the base period Stock distribution and migration is assumedconstantover timeand is representedby theaveragedistributionofCWTrecoveriesduringthebaseperiodHoweverdifferencesbe-tweentheabundancedistributionandmigrationpatternofstocksduringthebaseperiodandtheyearbeingevaluatedwilldecreasetheaccuracyof theestimatedstock-specificexploitation rates forthefisheryunderevaluationSignificantdeficienciesinFRAMmodelpredictionsforChinooksalmonfisheryassessmentduetotheuseofincompleteandoutdatedbaselinedatahaverecentlybeendemon-strated(MoranDazeyLaVoyampYoung2018)
Forcohosalmonutilityof theCWTsystemhasbeenerodedbytheextensivereleaseofadiposefin-clippedindividualswithoutCWTs and by the sharply declining number of CWTs recoveredfromrecentfisherysamplingThroughdirectqueryoftheRegionalMark Information System (RMPC 2018) it was determined thattherewere2664CWTs(averageof888CWTsannually)recoveredfrom commercial and recreational fisheries in BC between 2014and 2016 of which 1172 (391 annually) CWTswere from cohosalmonofCanadianoriginIncomparisontheaverageannualnum-berofCWTs recovered fromcoho salmonduring the seven-yearFRAMbaseperiodwas27119CWTsofwhich15436CWTswerefrom coho salmon of Canadian origin Thus the average annualnumberofCWTs recovered fromBC fisheriesduring2014ndash2016
K E Y W O R D S
coded-wiretagsCohosalmonfisherymanagementgeneticstockidentificationgenotypingbysequencingparentage-basedtagging
emspensp emsp | emsp3BEACHAM Et Al
was33andthoseofCanadianoriginwas25oftherespectiveannual numbers for the base period largely due to a substantialreductionincatchbutalsocoincidentwithreductioninthenum-berofCWTsappliedSinceestimationoffisheryimpactswiththeFRAM is dependent on CWT recovery information the greatlydecreased number of CWT recoveries in recent years will haveincreased the variance of the estimated stock-specific catch andexploitationratesinfisheries(Hinrichsenetal2016ReisenbichlerampHartmann1980)
Anewcost-effectiveapproachtotheassessmentandmanage-mentofwildcohosalmonandtheassociatedhatcheryproductionand fishery management is needed Anderson and Garza (2006)noted that parentage-based tagging (PBT) provides equivalent in-formation (hatcheryof releaseageof individual) forhatchery fishasdoCWTsimplementationofPBTthusmayovercomeproblemsassociatedwithCWT-basedassessmentandmanagementof cohosalmon fisheries inBCAdditionally PBTprovides ameansof im-provedhatcherybroodstockmanagementaswellasassessmentofhatchery-wild interactions in salmonids Unlike CWT-based man-agement PBT-informed hatchery and fishery management wouldbenefit from the complete adipose-clipping of hatchery-producedsalmonAsignificantadvantageofthecombinationofmassmarkingandPBT implementation is thecapabilityto identifyvisuallysam-ple and if desired remove hatchery fish of local and stray originin threatenedwildpopulationsMoreover PBTentails genotypingtheentirehatcherybroodstockandenablestheidentificationofallhatcheryprogenybyparentageassignment(Anderson2012Wang2016)thusenablingaldquomarkraterdquoofvirtually100ofhatcheryfishSteeleetal(2013)demonstratedtheequivalencyofCWTandPBTinaninitialevaluationofpopulationandageassignmentinsteelheadtrout(Oncorhynchus mykiss)oftheSnakeRiverbasinintheColumbiaRiverdrainageHessetal (2016)expandedtheapproachbyusingboth PBT and genetic stock identification (GSI) to investigate runtiming of steelhead trout in the upper Columbia River drainageTheseapplicationsconfirmedthecapabilityofacombinedPBT-GSItechnologytoprovideequivalentorbetter identificationoffishastheCWTmethodbutwerelimitedingeographicscaleInthisstudyweexaminewhether theexistingPBTandGSI systemof identifi-cationforCanadian-origincohosalmon(Beachametal2017)canprovidetheinformationrequiredforimprovedassessmentandman-agementofcohosalmonmixed-stockfisheriesinBritishColumbiacoveringamuchbroadergeographicrangeofcohopopulationsfromAlaskatoOregon
Although propermanagement of hatchery production and as-sociated fishery exploitation requires properly defined objectivesandmonitoring (Flagg2015) it is not currentlypossible toevalu-ate each Canadian enhancement project separately (TompkinsHamiltonBatemanampIrvine2016)ForcohosalmonCWTsarenotappliedtoreleasesfromsomeofthelargesthatcheriesinsouthernBC(ChilliwackRiverCapilanoRiverChehalisRiverConumaRiverNitinatRiverandTenderfootCreek)duetofundinglimitationsandthus theirspecificcontributionstohighlymixed-stockoceanfish-eriesareunknownInfactCWTsareappliedonlytocohosalmon
juvenilesreleasedfromsixFisheriesandOceansCanadalargepro-duction hatcheries three on the east coast of Vancouver Island(QuinsamRiverPuntledgeRiverandBigQualicumRiver)oneonthewestcoastofVancouverIsland(RobertsonCreek)oneinthelowerFraserRiverdrainage(InchCreek)andoneintheThompsonRiver(amajortributaryoftheFraserRiver)drainage(SpiusCreekwheretheSalmonRiverandEagleRiver juvenilesmaybe reared)CWTsarealsoappliedatsmallerfacilities(SeymourRivernearVancouverTobogganCreekandZymacordRiver intheSkeenaRiverdrainageinnorthernBC) and somenaturally spawned indexpopulations innorthernBC
The 43 coho salmon CUs originally identified by Holtby andCiruna(2007)havebeenmodifiedtoacurrentnumberof44CUsPrice Rosenberger Taylor and Stanford (2014) Price EnglishRosenbergerMacDuffeeandReynolds (2017)suggestedthatanysuitable assessment technique must provide individual resolutionfor all CUs to meet the conservation requirements of CanadarsquosWSPTherecanbepossible inconsistenciesbetweenCUsandex-istingfisherymanagementunits(MUs)(IrvineampFraser2008)withdifferencesbetweenCUsandMUschallengingmanagerswhoareresponsibleforbothassessingbiologicalstatusofCUsandfisheryobjectivesforMUs(HoltampIrvine2013)Giventhelimiteddistribu-tionofcohosalmonpopulationsmarkedwithCWTsinBCitisclearthatCWTscannotprovidetheCUresolutionrecommendedbyPriceetal(2014)Priceetal(2017)forwildpopulationassessmentnortheMUresolutionnotedbyHoltandIrvine(2013)forhatcheryandfisheryassessmentandmanagement
ThecurrentstudyisanevaluationoftheapplicationofthePBT-GSImethodologyoutlinedbyBeachametal(2017)tocohosalmonfisheriesinBCtodeterminewhetherthegenetictechnologiescanbe used to providemore information on fishery contributions byhatcheryandCUthanisavailablefromCWTsCommercialandrec-reationalcohosalmonfisheriesandriverescapementsforselectedpopulationsweresampledforbothCWTsandgenotypesWeeval-uatedthepopulation-levelresolutionobtainedfromCWTsandthegeneticmethodologybyCU for all 2017and some2016 fisheriesinwhichcohosalmonwerecaughtcatchestimationbyCUforthefisheriessampledandstock-specificexploitationrate forselectedpopulationsofcohosalmoninBCGenotypingbysequencingmeth-odologywasusedtogenotypecohosalmonat304singlenucleotidepolymorphisms(SNPs)in304ampliconsCompletebroodstockge-notypingforPBTanalysiswasconductedin2014for20hatchery-enhanced populations that included genotyping 6061 individuals(964genotypingsuccessrate)andastockidentificationbaselinecomprisingsome267populationsrangingfromsoutheastAlaskatoOregonwasemployedforGSIAcomparisonofthepopulation-spe-cificcontributionstomixed-stockfisheriescatchandexploitationratesestimatedwithCWTsandPBT-GSItechnologieswasmadeWeconcludethatageneticapproachcanemulateandimproveupontheresultsavailablefromthecurrentCWTprogramforassessmentandmanagementofcohosalmonenhancementandfisheriesinBCandprovidecriticalinformationtoimprovewildcohosalmonassessmentandconservation
4emsp |emsp emspensp BEACHAM Et Al
2emsp |emspMETHODS
21emsp|emspFishery sample collection
Atotalof8006 individualsweregenotyped fromfisherysamplescollected in2016and2017 In2016sampleswerecollectedfromadiposefin-clippedandunclippedcohosalmonlandedintherecrea-tionalfisheryintheStraitofGeorgiaJuandeFucaStraitandalongthewestcoastofVancouverIslandTheonlyexceptionswereArea21andArea121whereonlyclippedindividualsweresampledandAreas20-1to20-4inAugustwhereonlyclippedindividualsweresampled Sampleswerepooled for analysis asoutlined inTable1withthelocationsofstatisticalareasoutlinedbyhttpswwwpacdfo-mpogccafm-gpmaps-cartesareas-secteursindex-enghtmlAdditionalcohosalmonsamples(bothclippedandunclipped)wereobtained froma troll test fisheryoperatingnearBrooksPeninsulaoff thenorthwest coastofVancouver Island a gillnet test fisherytargetingsockeyesalmon(Oncorhynchus nerka)nearRoundIslandinnorthernJohnstoneStraitandacommercialdemonstrationfreezertrollfisheryinthecentralcoastofBC(Figure1)
In2017forthenorthern(AreaF)freezertrollfisheryselectedfreezerboats(28offleet)wererequiredtokeepheadsofallcohosalmoncaughtwiththemarktype(adiposefinclippedornot)un-knownforanindividualheadUponlandingtheheadswerecountedand checked electronically for CWTs and randomly sampled to amaximumof50headsperdeliveryIfaCWTwasdetectedtheheadwassenttoacentralCWTheadrecoverylaboratoryinVancouverBCwheretheDNAsamplewassubsequentlytakenIfaCWTwassuccessfully recoveredanddecoded froman individualhead from
commercialrecreationalorFirstNationsfisheriesandagenotypewassuccessfullyobtainedforthe individualsample thenthegen-otypesofallof these individualswerepooled intoasinglemixed-stocksampleofknownoriginandknownage inorder toevaluateaccuracyof stock compositionsbyCUandpopulationFieldDNAsamplesweretakenonlyfromindividualswithnoCWTsdetectedInthenortherniceboattrollfisherytheclipstatusofthefishinthecatchwasknownSamplesof clippedcohosalmonwereobtainedfrom this fishery as an ancillary aspect of standard Fisheries andOceansCanadacontractcatchsamplingforCWTsOnlyclippedin-dividualswereexaminedthroughthisprogramwithsimilarsamplingprotocolsasoutlinedinthefreezerboatsamplingInparticularonlyclippedfishnotcontainingaCWTweresampledinthefieldthroughthisprogramwithheadscontainingaCWTsenttotheheadrecov-ery laboratory inVancouverClipped fish constituted16of thecatchinthefisherylargelyduetothewildoriginofthecatchsoitwasimportanttoobtainsamplesfromunclippedindividualsaswellSamplesofunclipped individualswereobtainedby sampling land-ingsforamaximumof25individualspervesselwithamaximumof100 individuals sampled per day Samples from theCentralCoastfreezertrolldemonstrationfisherywereobtainedviaasimilarpro-tocolasoutlinedforthenorthernfreezertrollsamplingTheoriginofthesamplesfromtherecreationalfisheryinBCincludedvolun-taryheadrecoveriesofadiposefin-clippedcohosalmonfromrecre-ationalfisheriesinBCaswellassomecreelsamplingofrecreationalcatches Samples from the recreational fisherywerederived fromclippedindividualsbuttheymaynothavebeenmarkedwithaCWTwhendeliveredtotheCWTheadrecoverylaboratoryThussamplesin2017wereobtainedfromall individualsthatwouldberoutinely
Fishery Reporting region Statistical area
Recreational JohnstoneStrait(JST) A11toA13
StraitofGeorgia-north(SOG-n) A14toA16
StraitofGeorgia-south(SOG-s) A17toA18A19-7toA19-12A28A29
StraitofGeorgiaJuandeFucaStrait(SOGJDF)
A19-1toA19-6A20-5
WestcoastVancouverIslandJuandeFucaStrait(WCVIJDF)
A20(exceptA20-5)
SouthwestVancouverIsland(SWVI21121)
A21A121
SouthwestVancouverIsland-inshore(SWVI-inshore)
A23A24
SouthwestVancouverIsland-offshore(SWVI-offshore)
A123A124
NorthwestVancouverIsland-inshore(NWVI-inshore)
A25toA27
NorthwestVancouverIsland-offshore(NWVI-offshore)
A125toA127
Test BrooksPeninsula A27A127A126
RoundIsland A12
Troll CentralCoast A6A7A8
TA B L E 1 emspReportingregionandDFOstatisticalareaswithinregionsforamalgamationofcatchsamplesLocationsofstatisticalareasareoutlinedonhttpswwwpacdfo-mpogccafm-gpmaps-cartesareas-secteursindex-enghtml
emspensp emsp | emsp5BEACHAM Et Al
processed under the CWT recovery program for coho salmon inBCInadditionsampleswerecollectedfromarecreationalfishery(clippedandunclipped individuals) fromHaidaGwaiinearLangaraIslandarecreational fisherynearRivers Inlet (Area8clippedandunclippedindividuals)inthecentralcoastthepreviouslynotediceboat northern troll fishery and direct creel sampling (clipped andunclippedindividuals)ofrecreationalfisheriesinsouthernBC
Monthly catch composition estimates were determined forclipped individuals from various fisheries in BC Seasonal catchcomposition estimateswere determined by recording the numberofindividualgenotypesavailablefromthefisherysamplingineachmonth andweighting individualmonthly samples by the recordedcatch in themonth Individualswere randomlychosen formonthsinwhich the number of available genotypeswas in excess of thenumberrequiredfortheseasonalsampleForexampleif60oftheannualcatchwasrecordedinonemonthand200genotypeswereavailablefromthefisherythenallindividualssampledwouldbein-cludedintheseasonalsamplebutcontributionsofindividualsfromothermonthswerescaledrelativetothemonthlycontributiontothe
seasonalcatchIfthecatchinoneothermonthwas20oftheannualcatchand150sampleswereavailablethen200(2060)=67genotypeswererandomlychosenfromtheavailable150genotypesforinclusionintotheseasonalsample
InthenorthernBCtrollfisherysamplerselectronicallycheckedclippedindividualsforthepresenceofaCWTandheadsfromthoseindividualscontainingaCWTweresubsequentlysenttothecentrallaboratory for CWT recovery and tissue sampling for subsequentgenotypingTissuesamplesfromclippedindividualswithnoCWTsweredirectlyprovidedforgenotypingInordertoestimatemonthlyandseasonalstockcompositionofclippedindividualsinthisfisherywepooledthegenotypesobtainedfromdirectfisherysamplingofclippedindividualswiththosesubsequentlysenttothecentrallabo-ratoryforCWTrecoveryinordertoprovideanappropriatepoolofclippedindividualstoestimatemonthlyandseasonalstockcompo-sitionsofclippedindividualsinthefisheryIneachsampleanalyzedover all BC fisheries thenumberof individuals identified viaPBTrelative to thenumberofgenotypes in the samplewas tabulatedandsummarizedoversamplefisheryandseason
F I G U R E 1 emspMapindicatinggeographiclocationsforfisherysamplingand20populationsforwhichparentage-basedtaggingwasappliedinestimationofstockcomposition
6emsp |emsp emspensp BEACHAM Et Al
22emsp|emspEscapement sample collection
In2016adiposefin-clippedjacks(age2years)weresampledfromeightpopulationswith573individualscollectedandgenotypedsuc-cessfullyJacksweredefinedonthebasisofbodysizeduringsam-plecollectionwith jacksvisiblysmallerthanage3-yearspawnersPopulations sampled were as follows Robertson Creek QuinsamRiver Puntledge River Big Qualicum River Goldstream RiverChilliwackRiverStaveRiverand InchCreekTheobjectiveofthesamplingwas to obtain a sample of known origin and known agebaseduponknowncollectionlocationinordertoevaluateaccuracyofassignmentsof the jacks tobothCUandpopulationunder theassumption of no straying among populations for the individualssampled
In 2017 escapements (non-broodstock hatchery and river re-turns clipped individualsonly)were sampledwhere samplingwasfeasibleEscapementsamplingfrom13populationswasconductedand1692 individualsweregenotypedEscapement sampleswerenot available from all 20 populations forwhich itwas possible toassignindividualsinthenon-broodstockescapementsviaPBTTheobjectiveofthesamplingwastoevaluatestrayingratesamongpop-ulationsandforthosehatcheries(ChilliwackRiver)whereparentalgenotypeswereassignedtospecificreleasegroupsprovide infor-mationon the relative rates of returnof different release groupsAs relative ratesof return for the releasegroupswereconsideredancillarytothemainfocusofthestudynoresultsfromthesereturnswereincludedinthecurrentstudy
23emsp|emspExploitation rate
WeestimatedexploitationrateofadultcohosalmoninBCfisheriesviabothCWTsandgeneticsExploitationrateforapopulationwasdefinedasadultcatch(adultcatch-escapement)ForCWTstheob-servednumberofCWTswascorrectedbyldquono-pinrdquotaglossratesandwasexpandedbythepopulationrsquostag-specificmarkingratesummedovertagcodesandexpandedagainbythesamplingrateforthefish-eryinordertoestimatecatchofhatchery-originindividualsTheob-servednumberofCWTsinescapementsamplingwasexpandedinasimilarmanner inordertoestimatethehatcherycontributiontothe escapement For genetics the seasonal kept catchof adiposefin-clippedindividualsinafisherywasmultipliedbyaseasonalstockcompositionestimateinordertoestimatepopulation-specificcatchTheabundanceofhatchery-originescapementwascalculatedastheestimatedescapementmultipliedbytheproportionofadiposefin-clippedadultsobserved in theescapement If partof the juvenileproductionwasnotadiposefinclippeduponreleasefromthehatch-erythenthehatcherycontributionstobothcatchandescapementwereunderestimatedbythismethod
24emsp|emspGenetic stock identification baseline
The initial baseline was outlined by Beacham et al (2017) andconsisted of 20242 individuals from 117 populations with the
distributionofpopulationsrangingfromsoutheastAlaskatoPugetSoundinWashingtonStateThebaselinehasbeensubsequentlyex-panded to include 40774 individuals from267populations rang-ingfromsoutheastAlaskatoOregonTheprimaryexpansionofthebaselineincludedasurveyofadditionalpopulationsinsouthernBCcoastalWashingtontheColumbiaRiverdrainageandOregonThefull baseline is outlined in Supporting Information Table S1 withthe populations fromBC arrangedbyCU andwithUnited StatesofAmerica(US)populationsarrangedbygeographic(reporting)re-gionBeachametal (2017)hadpreviouslydemonstratedastrongregionalpopulationstructureandreportedthatoverall117popula-tionsaverageself-assignmentaccuracytopopulation(weightedbypopulation sample size)was 855 for individualswith an assign-mentprobabilitylt050excludedand928forindividualswithanassignmentprobabilitylt085excludedHighlevelsofaccuracyforGSI assignment of individuals to specific populations when com-binedwithassignmentsviaPBTprovideapowerful technique forassignmentofindividualsofunknownorigintosomepopulations
25emsp|emspLibrary preparation and genotyping
ThedetailedprocedureforlibrarypreparationandgenotypingwasoutlinedbyBeachametal(2017)SummarizedbrieflyDNAwasex-tractedfromcohosalmontissuesamplesviaaChelexextractionandtheDNAconcentration normalized to40ngμlwith a Tecan LiHarobotApanelofprimersdesignedtoamplifyspecificsegmentsofDNAthatcontainedSNPsofinterestwasappliedtotheextractedDNAwithprimersequencesforeachampliconoutlinedbyBeachametal (2017SupportingInformationTableS2)TheinitialmultiplexPCR amplification of 304 target ampliconswas conductedwith acocktail of 2μl of normalizedDNA extract 5μl of 2X Ion Agriseqprimerpool2μlofIonAgriseqHiFimixand1μlofddH2O Thermal cyclingwasconductedin96-wellPCRplates(oneindividualperwell)withthefollowingconditionsforPCR99degCndash2min17cycles(99degCndash15s60degCndash4min)10degChold
FollowingtheinitialPCRasecondstepemployingathermalcy-clerwasconducted thatpartiallydigested theprimerson theam-pliconsandthereactionconductedwiththefollowingconditions50degCndash10min55degCndash10min60degCndash20min10degCholdA thirdandfinalstepemployingathermalcyclerwasinitiatedtoligatethebarcodes(768individualcodes)totheampliconsandwasconductedwiththefollowingconditions22degCndash30min70degCndash10min10degCholdLibrarieswerepurifiedbyadditionof225μlofAgencourtregAMPureregXPmagneticbeadstoeachlibrarytheplatewasplacedonamagnetic racksupernatantdiscardedandthebeadswashedtwice in70ethanolThepurified librarieswerethenelutedwith25 μloflowTEand20μlofthesupernatanttransferredtoafresh96-well trayNext eachof the768prepared librarieswaspooledintoasingletubeforprocessingontheIonChefreg(ThermoFisherScientific)Twotubesofpooled librarieswereprocessedconsecu-tivelyontheIonChefandthus1536individualswereprocessedon a single run of the Ion ChefOne tube of the pooled librarieswasloadedontoeachP1regchipv3andthusampliconsfrom768
emspensp emsp | emsp7BEACHAM Et Al
individualsweredistributedoneachP1chipwith1536individualsprocessedbetweentwochipsThechipswerethenloadedontotheIonTorrentProtonsequencerAfter thesequencingrunwascom-pletedcomparisonswiththereferencegenomeoftherainbowtrout(O mykiss)(Berthelotetal2014)supplementedwiththesequencescontaining theobservedcohosalmonSNPswereconductedwithProton softwareVariant Callerreg and SNP genotypes at the sitesspecified by the hotspot filewithin target regionswere called byVariantCallerThehotspotfilecontained304SNPsiteswithoneSNPscoredateachampliconGenotypesatallavailableSNPsforanindividualwereassembled toprovideamulti-locus individualgen-otype The species identification SNPOkiOts_120255‐113 (StarksClementoampGarza2016)andsex identificationSNPOts_SEXY3‐1 wereomittedfromsubsequentparentageandGSIanalysesleaving302SNPs
26emsp|emspIdentification of individuals
AsnotedpreviouslyPBTandGSIwereusedconcurrently toesti-mate fishery stock composition Initially PBT was used and theanalysiswas conductedwhere the genotypes of individuals to beidentifiedwerematched to the genotypes of prospective parents(COLONY JonesampWang 2010Wang 2016) If all individuals ina hatchery broodstock are sampled and subsequently genotypedthenalloffspringfromthebroodstockaregeneticallymarkedThegenotypes of individuals of unknown origin are statistically com-pared with the genotypes of potential parents and if a match ismadetheoffspringareassignedtotheparentsandthustheori-ginandageoftheindividualsaredeterminedParentageassignmentsoftwarewasutilizedtoassignoffspringtoparentsasCOLONYcanproduceassignmentswhen thegenotypeofoneof theparents ismissingeitherduetoamissingparentalsampleor failure topro-duceaparentalgenotypefromanexistingsampleGiventhatPBTassignments for20potential populationswereevaluated for eachfisherysampleCOLONYwasrunwithallbroodstocksampleddur-ing2014inputasasingleunitforanalysisoffisherysampleswithnodifferentiationamongpopulationsAlthoughtheCOLONYassump-tionofasinglepopulationintheparentpoolwasviolatedanalysisofknown-originsamplesindicatedthatveryhighlevelsofaccuracywereachievedinassignmentswhenpoolingofpotentialparentsincontributing populationswas conducted Two-parent assignmentswere accepted only when both assigned parents originated fromthesamepopulationotherwisetheindividualwaspassedtogeneticstockidentification(GSI)forpotentialassignmentTwo-parentandsingle-parentassignmentswereacceptedonlywhentheprobabilityofcorrectassignmentwasge085fortheparentpairotherwisetheindividualwaspassedtopotentialassignmentbyGSIAdditionallyforsingle-parentassignments tobeaccepted thePBTassignmenttopopulationhad tobepart of theCUassigned to the individualviaGSI Individuals forwhichnoprospectiveparentswere identi-fiedinthebroodstockwerepassedtoGSIforpotentialassignmentPolygamousmatingwasassumedfortheCOLONYanalysisSimplepairwisecomparisonsbetweenoffspringandpotentialparentswere
conductedThebaselineforindividualssampledinthe2016escape-ments (jacks)and2017fisheries includedallbroodstockssampledin2014as these individualsarepredominately threeyearsofage(Sandercock1991)Jacks inthe2017escapementwere identifiedvia body size and subsequent assignment to parents in the 2015hatcherybroodstocksIndividualswithmorethan120missinggeno-typeswereeliminatedfromfurtheranalysesAnestimatedgenotyp-ingerrorrateof1wasusedforCOLONYassignmentsPreviouslyBeachametal(2017)hadreportedthatanaveragegenotypingerrorrate of 107 (1220discrepancies in 114105 comparisons) or analleleerrorrateof053(1220discrepanciesin228210compari-sons)wasobservedoverthe302SNPsscoredTheparentpairout-putfilewasthebasicfileusedinsubsequentanalyses
Thesecondmethodof individual identification isGSI inwhichthegeneticprofilesofwholepopulationspotentiallycontributingtoamixed-stocksampleareusedtoestimatetheoriginofeachindivid-ualinthesample(RUBIASMoranampAnderson2018)ThisanalysiswasrestrictedtothoseindividualsunassignedviaCOLONYForeachsample individuals not assigned by COLONYwere then assignedwithRUBIASThepopulationposteriormeansfilewasthebasicfileusedforsubsequentanalysesandthisfilecontainedtheprobabilityofassignmentoftheindividualtoeachofthe267populationsinthebaselineStockcompositionwasestimatedthroughthecombinationoffilesgeneratedwithbothCOLONYandRUBIAS Individualsas-signedviaCOLONYwereassignedaprobabilityof100oforiginat-ingfromtheidentifiedpopulationwitha000probabilityassignedtoallotherpopulationsinthebaselineThislevelofassignmentac-curacyviaPBTwasobservedpreviouslyforcohosalmonwiththepanelofSNPsemployedinthecurrentstudy(Beachametal2017)ThesedatawerethencombinedwiththeprobabilityofassignmentforthoseindividualsunassignedviaCOLONYtoeachofthepopu-lationsinthebaselineviaRUBIASAfteraninitialburn-inof25000iterations the last1000 iterations from theMonteCarloMarkovchainfromRUBIASwereusedtoestimatetheoriginofindividualsandstockcompositionwiththemeanallocationtoeachpopulationinthebaselinedeterminedStandarddeviationsofestimatedstockcompositionswere alsodetermined from the last1000 iterationsfrom theMonteCarloMarkovchain Stock compositionbyCUorreportinggroupwasdeterminedbysummationofallocationstoallpopulations in the baseline that belonged to the CU or reportinggroupunderconsideration
3emsp |emspRESULTS
31emsp|emspEstimation of stock composition for known‐origin samples
Genotypeswere available from573 jacks sampled in2016acrossthreeCUs and eight populations in southernBC Estimated stockcompositionbyCUand reportinggroup for this combinedsamplewas accuratewith an error utilizingonlyGSI of le04byCU forthreeCUspresentinthesample(Table2)WithbothPBTandGSIutilized theaverageerrordeclinedtole02byCUfor threeCUs
8emsp |emsp emspensp BEACHAM Et Al
TAB
LE 2
emspAccuracyofregional(UnitedStates)andconservationunit(Canada)estimatedstockcomposition()foramixed-stocksampleof573known-origincohosalmonjackssampledin
2016and620coded-wiretaggedindividualssampledin2017fisheriesinBCStandarddeviationoftheestimateisinparentheses
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
SoutheastAlaska
00
00(00)
00(00)
90
68(11)
68(11)
AlsekRiver
00
00(00)
00(00)
00
00(00)
00(00)
LowerStikine
00
00(00)
00(00)
00
04(03)
04(03)
LowerNass
00
00(00)
00(00)
63
64(10)
62(10)
UpperNass
00
00(00)
00(00)
00
04(03)
04(04)
PortlandSound-Observatory
Inlet-PortlandCanal
00
00(00)
00(00)
00
00(00)
00(00)
SkeenaEstuary
00
00(00)
00(00)
00
00(00)
00(00)
LowerSkeena
00
00(00)
00(00)
16
12(05)
12(05)
MiddleSkeena
00
00(00)
00(00)
69
39(08)
39(09)
UpperSkeena
00
00(00)
00(00)
06
40(10)
40(09)
HaidaGwaiindashGrahamIslandLowlands
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-East
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-W
est
00
00(00)
00(00)
00
00(00)
00(00)
NorthernCoastalStreams
00
00(00)
00(00)
00
00(00)
00(01)
HecateStraitMainland
00
00(00)
00(00)
00
07(05)
06(04)
MusselndashKynoch
00
00(00)
00(00)
00
00(00)
00(00)
DouglasChannelndashKitimatArm
00
00(01)
00(01)
00
00(01)
00(01)
BellaCoolandashDeanRivers
00
00(00)
00(00)
00
00(00)
00(00)
RiversInlet
00
00(01)
00(00)
00
00(00)
00(00)
SmithInlet
00
00(00)
00(00)
00
00(00)
00(00)
SouthernCoastalStreamsndashQueen
CharlotteStraitndashJohnstoneStraitndash
SouthernFjords
00
00(01)
00(01)
00
00(02)
00(02)
HomathkondashKlinakliniRivers
00
00(00)
00(00)
00
00(00)
00(00)
GeorgiaStraitMainland
00
00(00)
00(00)
00
00(00)
00(00)
HoweSoundndashBurrardInlet
00
00(00)
00(00)
06
00(01)
00(00)
EastVancouverIslandndashGeorgiaStrait
351
345(20)
348(20)
131
134(14)
140(14)
EastVancouverIslandndashJohnstone
StraitndashSouthernFjords
00
00(00)
00(00)
00
00(01)
00(00)
NahwittiLowland
00
00(01)
00(00)
18
17(06)
17(06) (C
ontin
ues)
emspensp emsp | emsp9BEACHAM Et Al
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
WestVancouverIsland
115
115(14)
115(13)
135
134(14)
135(14)
Clayoquot
00
00(00)
00(00)
00
02(03)
02(02)
JuandeFucandashPachena
00
00(01)
00(01)
00
00(00)
00(00)
LowerFraser
534
539(21)
536(21)
185
187(15)
188(15)
Lillooet
00
00(00)
00(00)
00
03(02)
02(01)
FraserCanyon
00
00(00)
00(00)
00
00(00)
00(00)
InteriorFraser
00
00(00)
00(00)
00
00(00)
00(00)
LowerThompson
00
00(00)
00(00)
08
07(03)
08(04)
NorthThompson
00
00(00)
00(01)
00
00(00)
00(00)
SouthThompson
00
00(00)
00(00)
05
06(03)
05(03)
Boun
dary
Bay
00
00(00)
00(00)
00
00(00)
00(00)
NooksackRiver
00
01(02)
01(02)
26
09(09)
10(08)
SkagitRiver
00
00(01)
00(01)
05
23(08)
22(08)
NorthernPugetSound
00
00(00)
00(00)
39
58(12)
55(13)
Mid-PugetSound
00
00(00)
00(00)
52
62(11)
63(11)
SouthernPugetSound
00
00(00)
00(00)
10
09(10)
06(08)
JuandeFucaStrait
00
00(00)
00(00)
00
02(03)
01(03)
HoodCanal
00
00(00)
00(00)
44
16(05)
16(06)
CoastalWashington
00
00(00)
00(00)
60
59(10)
59(10)
ColumbiaRiver
00
00(00)
00(00)
32
32(07)
32(07)
Oregon
00
00(00)
00(00)
00
00(01)
00(00)
TAB
LE 2
(Continued)
10emsp |emsp emspensp BEACHAM Et Al
presentinthesample(Table2)Theaverageerrorfor45CUsorre-portinggroupsabsentinthesamplewas00bybothGSIandPBT-GSIBypopulation543(948)ofthejackswereassignedviaPBTwith 100 accuracywith respect to population of origin and age(Table3)Estimatedstockcompositionfor20populationswhereitwaspossibletousebothPBTandGSIinestimationofstockcompo-sitionwasaccurateFortheeightpopulationspresentinthesampleofjackstheaverageerrorutilizingGSIonlywas05perpopulationdecliningto02utilizingbothPBTandGSIundertheassumptionthatthecollectionlocationwasanaccuratereflectionofjackoriginTherewasanaverage00errorforthe12populationswithnorep-resentation inthesamplewithutilizationofeitherGSIorPBT-GSI(Table4)ThelargesterrorutilizingbothPBTandGSI(underestima-tionby06ofactualvalue)wasobservedintheGoldstreamRiverpopulationwhichalsodisplayedthe lowestpercentage (704)ofjacksidentifiedviaPBT(Table3)PBTwhencombinedwithGSIpro-ducedmoreaccurateestimatesofCU-specificandpopulation-spe-cificstockcompositionthanavailablewithonlyGSI
Of the 3054 adipose fin-clipped coho salmon samples re-ceivedforpotentialgenotypingfromacentrallaboratorywhere2017fisherysampleswereprocessedforpotentialCWTrecov-erygenotypeswereobtained for2533 (829)of the individ-uals processed There were 750 CWTs recovered from 3054headsor snouts examined and genotypeswereobtained from622 (829) of the individuals that carried a CWT GenotypesofcohosalmonwithoutCWTsbutsentfromtheCWTheadre-covery laboratorywereobtainedfrom1911of2304 individu-als (829) Release information associatedwith theCWTwasavailable for 620 of the 622CWTs associatedwith genotypedindividuals and these 620 individuals constituted a sample ofknown origin There were individuals from 20 regions or CUspresentinthissampleandforthese20regionsorCUstheav-erageerrorofestimationutilizingonlyGSIwas09perregionorCUand09forbothPBTandGSIcombined(Table2)CWTswererecoveredoriginatingfromsevenpopulationswhereitwaspossibletoincorporatebothGSIandPBTinestimationofstockcompositionFor these sevenpopulations theaverageerror inestimatedstockcompositionsperpopulationwas20withGSIand 03 with PBT-GSI (Table 4) There was an average 00errorforthe13populationswithnorepresentationinthesam-ple with utilization of either GSI or PBT-GSI (Table 4) Similartothesampleof jacksPBTwhencombinedwithGSIproducedmore accurate estimates of population-specific stock composi-tionthanavailablewithonlyGSI
32emsp|emspEvaluation of 2016 fishery sampling
Asbroodstocksamplingatselectedhatcheries insouthernBCdidnot commence until 2014 it was not possible to obtain PBT as-signments for any fishery samples collected in 2016 HoweverGSI analysis of individuals derived from recreational fishery sam-ples intheStraitofGeorgiaJuandeFucaStraitandoffthewestcoastofVancouverIslandinsouthernBCindicatedthattherewas
virtually no contribution from regions orCUsnorthofVancouverIsland (Supporting Information Table S2) The only exceptionwasin thenorthwestcoastofVancouver Islandboth inshoreandoff-shorewhereupto12ofthesamplewasidentifiedasoriginatingfromnorthernandcentralBCCUsUS-origincohosalmonwerees-timatedtohavecontributedapproximately70ofsummerrecrea-tionalsamplesderivedfromoffshorefisheriesalongthewestcoastofVancouverIslandaswellasintheentrancetoJuandeFucaStrait(A21A121)InmoreinshoreareasalongthewestcoastofVancouverIslandcohosalmonfromlocalCUswerethedominantcontributorstothefisheryForexamplethewestcoastofVancouverIslandCUcontributedapproximately80ofthe39-individualAugustsamplefromtheinshoresouthwesternVancouverIslandregionIntheStraitofGeorgia localCUswereagainthemaincontributorstotherec-reationalfisheryduringthesummerwiththeHoweSoundndashBurrardInletCUcontributingupto40insomesamplestheeastcoastofVancouverIslandCUuptoapproximately30andthelowerFraserRiverCUuptoapproximately40WhilethelowerFraserRiverCUwasanimportantcontributortotherecreationalfisheryintheStraitofGeorgiaandJuandeFucaStraititscontributiontofisheriesoffthewestcoastofVancouverIslandwaslimitedTheNooksackRiverregion innorthernWashingtonStatewasalso identifiedasacon-sistent contributor to recreational fishery samples in the Strait ofGeorgia insomecasescontributingupto20ofthesampleana-lyzed(SupportingInformationTableS2)
StockcompositionswereavailablefromtwotestfisheriesoneatrolltestfisherynearBrooksPeninsulabynorthwesternVancouverIslandandtheotheragillnettestfisheryinupperJohnstoneStraitIn the Brooks Peninsula fishery Vancouver IslandCUs comprised40ndash60ofthesampleswithaminorcontributionofFraserRiverCUs (lt10) The five interior Fraser River CUs of conservationconcern comprised00ndash33of the sampleswhereas themoreabundant lower Fraser River CUs comprised only 00ndash55 ofthesamples Individuals fromFraserRiverCUswerenotdetectedduringJulyinArea127theoffshoreportionofnorthernnorthwest-ernVancouverIslandbuttheyweredetectedinArea126themoresoutherlyoffshoreportionofnorthwesternVancouverIslandIntheJohnstoneStrait(RoundIsland)testfisheryindividualsfromFraserRiverCUswerescarcebutthefiveinteriorFraserRiverCUscom-prised30ndash35of the sampleswhereas the lowerFraserRiverCUscomprisedonly00ndash65ofthesamplesThemajorcontrib-utors to the samplesoriginated from theeast coastofVancouverIsland(24ndash45)andHoweSoundndashBurrardInletCUs(9ndash11)aswellascoastalWashington(8ndash15)(SupportingInformationTableS2)
ThefinalfisherysampledwasademonstrationcommercialtrollfisheryincentralcoastalBC(A6andA7)ContributionsfromlocalCUswerehigherinsamplesfrommoreinshoreareas(A7-7August212southernCUsandregions)withcontributionsfromsouth-ernCUshigherinthemoreseawardareas(A6-9September862southernCUsandregions)LikethewestcoastofVancouverIslandmigrating stocks were more likely to be found in more seawardlocations
emspensp emsp | emsp11BEACHAM Et Al
33emsp|emspEvaluation of 2017 fishery sampling
Onesourceoffisherysampleswasdirectfisherysamplingofindi-viduals(bothadiposefin-clippedandunclipped)wherethefocuswas not associated with potential CWT recovery In this casesamplingofunclippedindividualsfromthenorthern(AreaF)trollfishery indicated that initially (July 10ndash12) northern and centralBCCUscontributed23tothecatchsouthernBCCUs53andsouthern US regions 23 (Supporting Information Table S3)Stock composition changed markedly in samples collected July13ndash18 with Alaska contributing 12 northern and central BCCUs contributing 72 and southern BCCUs contributing 16withnosouthernUScontributionSamplescollectedduringJuly22ndash28revealedadominantcontributionfromnorthernandcen-tralBCCUs(96)andCUsfromthisregioncontributedbyfarthelargestportionof the catch in subsequent sampleswith south-ernBCCUsandsouthernUSregionslargelyabsent(SupportingInformation Table S3) The average contribution to the weeklysamples from July 22 onwardswas 17 southeast Alaska 16HaidaGwaiindashGrahamIslandLowlandsCU15DouglasChannelndashKitimatArmCU 12Hecate StraitMainlandCU 8NorthernCoastalStreamsCUand7LowerNassCUNotunexpectedlysampling of only clipped individuals from the fishery indicatedverysubstantialdifferencesinstockcompositionbetweenclippedandunclippedindividualsTheaveragecontributiontotheweeklysamples of clipped individuals from July 19 onwards was 32coastalWashington11mid-PugetSound10westVancouverIsland CU and 5 for each of northern Puget Sound and Juande Fuca Strait (Supporting Information Table S4) However theobservedcliprateinthefisherywas16sothecatchwasdomi-natedbyunclippedindividualsTheSeptember2ndash8samplefromthe Haida Gwaii recreational fishery near Langara displayed astockcompositionof42HaidaGwaiindashGrahamIslandLowlandsCU15HecateStraitMainlandCU10LowerSkeenaCUand8 Northern Coastal Streams CU similar to that observed inthe troll fisherywith the local CU (HaidaGwaiindashGraham IslandLowlands) inhigherproportionsaswouldbeexpectedinamoreinshorefishery(SupportingInformationTableS3)
Directsamplingof thecentralcoast troll fisheryrevealedthatonaverage theDouglasChannelndashKitimatArmCUwasthe largestcontributortothefishery(24)followedbytheNorthernCoastalStreamsCU(19)BellaCoolandashDeanRivers(18)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(10) (Supporting InformationTableS3)withallCUsproximaltothelocationofthefisherySamplesfromtherecreationalcentralcoastfisheryindicatedthattheBellaCoolandashDeanRiversCUwasthedominant contributor to the fishery (32) followedby theRiversInletCU(19)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(15)andDouglasChannelndashKitimatArmCU(7) (SupportingInformationTableS3)GiventhelocationofthefisherydominantcontributionsfromtheseCUsweretobeexpected
AsnotedpreviouslysampleswerealsoavailablefromvirtuallyallindividualssampledfromcommercialrecreationalandFirstNationsfisheriesthatwouldberoutinelyprocessedforCWTrecoveryinayearbyacentrallaboratoryIndividualheadsfromcommercialfish-eries sent to the central laboratoryhadbeenpreviously screenedelectronically for thepresenceofaCWTVoluntaryheadsamplesfromadiposefin-clippedindividualsfromrecreationalfisheriesweresenttothecentrallaboratorybutmanyoftheseindividualsdidnotcontainaCWTastherewasnopriorscreeningof these individu-als Stock composition of clipped individuals in the northern trollfishery has been previously outlined above Stock composition inthenorthern recreational fishery indicatedawidegeographicdis-tributionofcontributions rangingfromcoastalWashington (16)thewestcoastofVancouverIslandCU(10)theupperSkeenaCU(9)southeastAlaskaregion(8)mid-PugetSound(7)andeastcoast of Vancouver IslandndashStrait of Georgia CU (6) (SupportingInformation Table S4) Stock composition of clipped individuals inthecentralcoastrecreationalfisherywasdominatedbytheHoweSoundndashBurrardInletCU(27)andeastVancouverIslandndashStraitofGeorgiaCU(25)(SupportingInformationTableS4)butgiventhelowobservedclip rate in the fishery (014) therewasvery littlecatchthatcouldbeidentifiedashatcheryorigin
Directsamplingofrecreationalfisheries insouthernBCwhichincluded both clipped and unclipped individuals in the sample
Population Genotyped Assigned assignment
accuracy of population assignment
Robertson 66 66 1000 1000
Quinsam 69 68 986 1000
Puntledge 21 19 905 1000
BigQualicum 84 84 1000 1000
Goldstream 27 19 704 1000
Chilliwack 195 183 938 1000
Inch 91 85 934 1000
Stave 20 19 950 1000
Total 573 543 948 1000
TA B L E 3 emspPercentageofadiposefin-clippedjacks(age2years)sampledin2016andassignedviaparentageanalysistoparentsin2014broodstockpopulationsPopulationofsamplingwasassumedtobethepopulationoforiginofthejacksforevaluationofaccuracyofassignment
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
2emsp |emsp emspensp BEACHAM Et Al
1emsp |emspINTRODUC TION
WildPacific salmon includingCoho salmonOnchorynchus kisutchhavebeensupplementedwithhatcherypropagationforoverhalfacenturyinsupportofincreasedoceanharvestandconservationofthreatenedpopulationsThepotentialnegativeeffectsofhatcheryproduction and the associated exploitation pressure that may beappliedtonaturalpopulationshavelongbeenrecognizedandhavegainedprominenceinrecentyears(ArakiBerejikianFordampBlouin2008Hilborn1992JonesCornwellBottomSteinampAnlauf-Dunn2018McClureetal2008)leadingtocallsforincreasedresponsi-bilityinmanagementofhatcheryproductionandofthemixed-stockfisheriesitsupports(Flagg2015HSRG2014)InCanadatheWildSalmon Policy (WSP) for Pacific salmonwas establishedwith thegoalofmaintainingandrestoringhealthyanddiversePacificsalmonpopulations making conservation of wild salmon and their habi-tatsthehighestpriorityforresourcemanagementdecision-making(FisheriesandOceansCanada2005)Fisheriesandhatcherysupple-mentation (termedenhancement inCanada)are tobemanaged insuchawayastoensurethatwildpopulationsaresafeguardedandharvestbenefitsaresustainableWildsalmonpopulationsareiden-tifiedandmaintained inconservationunits (CUs) that reflect theirgeographicecologicalandgeneticdiversity
CohosalmonarecaughtincommercialrecreationalandFirstNations fisheries in British Columbia and determination of theimpact of these fisheries is of fundamental importance to statusassessmentforwildpopulationsofconservationconcernandman-agementoflarge-scalehatcheryproductionCurrentandhistoricalassessmentoffisheriesimpactshasbeenconductedwiththeap-plicationofcoded-wiretags(CWTsJeffertsBergmannampFiscus1963)CWTsareappliedtojuvenilefishpriortotheirhatcheryre-leaseandrecoveredfromadultfishheadscollectedfromfisherieshatchery broodstocks and in-river escapement sampling OncerecoveredthetagsaredecodedtodeterminethehatcheryoriginandageoftheindividualfishOriginallyonlycohosalmonmarkedwithaCWTalsoreceivedanadiposefinclippriortohatcheryre-leasewith theexternally visible clipmarkallowingCWT-markedfish to be identified visually and sampled from fisheries or rivercollections
Sincethelate1990sallcohosalmonreleasedfrommanyhatch-eries in southern British Columbia (BC)Washington andOregonhave received an adipose fin clip (termedmassmarking) in orderto facilitatemark-selective fisheries intended to harvest hatcherysalmononlywithmost clipped individuals carrying noCWTThisapproachhasresultedinreducedexploitationofnaturallyspawnedcohosalmonespeciallyinsportfisheriesbutthepresenceofmanyadipose-clippedsalmonwithoutaCWThasimpairedtheefficiency
ofCWT recovery In spite of implementation of an electronic tagdetectionsystemtopre-screenaportionofthecommercialcatchtoidentifysalmonwithaCWTtheprocessingofmanyheadswithouta CWT from voluntary recreational recoveries and the increasingcostsofCWTapplicationandrecoveryhavecauseddegradationofthe informationobtained from the currentCanadian coho salmonassessmentprogram
For coho salmon fisheries impacts are evaluated through theFishery Regulation AssessmentModel (FRAM PFMC 2008) TheFRAMisadiscrete-time-stepage-structuredcomputermodelusedtopredicttheimpactsofavarietyofproposedfisheryregulationsforasinglemanagementyearinimplementationofprovisionsofthePacificSalmonTreaty (PST) related tocohosalmonCanadianandAmericanassessmentstaffusethebilaterallyagreeduponFRAMtoestimatefisheryimpactsPriortoeachfishingseasonbothcountriesincorporatestock-specificconservationconstraintsidentifiedinthecohosalmonchapterofthePST intoFRAMevaluationofplannedfisheriesApplicationoftheFRAMrequiresuseofdataderivedfromfisheryandescapementrecoveriesofCWTsandterminalarearunsizeestimatesforreturnyearsfromabaseperiod(1986ndash1992)Thecurrentexpectedcatchofaspecificstockisestimatedastheprod-uctof(theexpectedabundanceofthestockinafishery)(thebaseperiodexploitation rate)(a correction factor) that relates theex-pectedcatchoreffortinaparticularyearrelativetothatobservedin the base period Stock distribution and migration is assumedconstantover timeand is representedby theaveragedistributionofCWTrecoveriesduringthebaseperiodHoweverdifferencesbe-tweentheabundancedistributionandmigrationpatternofstocksduringthebaseperiodandtheyearbeingevaluatedwilldecreasetheaccuracyof theestimatedstock-specificexploitation rates forthefisheryunderevaluationSignificantdeficienciesinFRAMmodelpredictionsforChinooksalmonfisheryassessmentduetotheuseofincompleteandoutdatedbaselinedatahaverecentlybeendemon-strated(MoranDazeyLaVoyampYoung2018)
Forcohosalmonutilityof theCWTsystemhasbeenerodedbytheextensivereleaseofadiposefin-clippedindividualswithoutCWTs and by the sharply declining number of CWTs recoveredfromrecentfisherysamplingThroughdirectqueryoftheRegionalMark Information System (RMPC 2018) it was determined thattherewere2664CWTs(averageof888CWTsannually)recoveredfrom commercial and recreational fisheries in BC between 2014and 2016 of which 1172 (391 annually) CWTswere from cohosalmonofCanadianoriginIncomparisontheaverageannualnum-berofCWTs recovered fromcoho salmonduring the seven-yearFRAMbaseperiodwas27119CWTsofwhich15436CWTswerefrom coho salmon of Canadian origin Thus the average annualnumberofCWTs recovered fromBC fisheriesduring2014ndash2016
K E Y W O R D S
coded-wiretagsCohosalmonfisherymanagementgeneticstockidentificationgenotypingbysequencingparentage-basedtagging
emspensp emsp | emsp3BEACHAM Et Al
was33andthoseofCanadianoriginwas25oftherespectiveannual numbers for the base period largely due to a substantialreductionincatchbutalsocoincidentwithreductioninthenum-berofCWTsappliedSinceestimationoffisheryimpactswiththeFRAM is dependent on CWT recovery information the greatlydecreased number of CWT recoveries in recent years will haveincreased the variance of the estimated stock-specific catch andexploitationratesinfisheries(Hinrichsenetal2016ReisenbichlerampHartmann1980)
Anewcost-effectiveapproachtotheassessmentandmanage-mentofwildcohosalmonandtheassociatedhatcheryproductionand fishery management is needed Anderson and Garza (2006)noted that parentage-based tagging (PBT) provides equivalent in-formation (hatcheryof releaseageof individual) forhatchery fishasdoCWTsimplementationofPBTthusmayovercomeproblemsassociatedwithCWT-basedassessmentandmanagementof cohosalmon fisheries inBCAdditionally PBTprovides ameansof im-provedhatcherybroodstockmanagementaswellasassessmentofhatchery-wild interactions in salmonids Unlike CWT-based man-agement PBT-informed hatchery and fishery management wouldbenefit from the complete adipose-clipping of hatchery-producedsalmonAsignificantadvantageofthecombinationofmassmarkingandPBT implementation is thecapabilityto identifyvisuallysam-ple and if desired remove hatchery fish of local and stray originin threatenedwildpopulationsMoreover PBTentails genotypingtheentirehatcherybroodstockandenablestheidentificationofallhatcheryprogenybyparentageassignment(Anderson2012Wang2016)thusenablingaldquomarkraterdquoofvirtually100ofhatcheryfishSteeleetal(2013)demonstratedtheequivalencyofCWTandPBTinaninitialevaluationofpopulationandageassignmentinsteelheadtrout(Oncorhynchus mykiss)oftheSnakeRiverbasinintheColumbiaRiverdrainageHessetal (2016)expandedtheapproachbyusingboth PBT and genetic stock identification (GSI) to investigate runtiming of steelhead trout in the upper Columbia River drainageTheseapplicationsconfirmedthecapabilityofacombinedPBT-GSItechnologytoprovideequivalentorbetter identificationoffishastheCWTmethodbutwerelimitedingeographicscaleInthisstudyweexaminewhether theexistingPBTandGSI systemof identifi-cationforCanadian-origincohosalmon(Beachametal2017)canprovidetheinformationrequiredforimprovedassessmentandman-agementofcohosalmonmixed-stockfisheriesinBritishColumbiacoveringamuchbroadergeographicrangeofcohopopulationsfromAlaskatoOregon
Although propermanagement of hatchery production and as-sociated fishery exploitation requires properly defined objectivesandmonitoring (Flagg2015) it is not currentlypossible toevalu-ate each Canadian enhancement project separately (TompkinsHamiltonBatemanampIrvine2016)ForcohosalmonCWTsarenotappliedtoreleasesfromsomeofthelargesthatcheriesinsouthernBC(ChilliwackRiverCapilanoRiverChehalisRiverConumaRiverNitinatRiverandTenderfootCreek)duetofundinglimitationsandthus theirspecificcontributionstohighlymixed-stockoceanfish-eriesareunknownInfactCWTsareappliedonlytocohosalmon
juvenilesreleasedfromsixFisheriesandOceansCanadalargepro-duction hatcheries three on the east coast of Vancouver Island(QuinsamRiverPuntledgeRiverandBigQualicumRiver)oneonthewestcoastofVancouverIsland(RobertsonCreek)oneinthelowerFraserRiverdrainage(InchCreek)andoneintheThompsonRiver(amajortributaryoftheFraserRiver)drainage(SpiusCreekwheretheSalmonRiverandEagleRiver juvenilesmaybe reared)CWTsarealsoappliedatsmallerfacilities(SeymourRivernearVancouverTobogganCreekandZymacordRiver intheSkeenaRiverdrainageinnorthernBC) and somenaturally spawned indexpopulations innorthernBC
The 43 coho salmon CUs originally identified by Holtby andCiruna(2007)havebeenmodifiedtoacurrentnumberof44CUsPrice Rosenberger Taylor and Stanford (2014) Price EnglishRosenbergerMacDuffeeandReynolds (2017)suggestedthatanysuitable assessment technique must provide individual resolutionfor all CUs to meet the conservation requirements of CanadarsquosWSPTherecanbepossible inconsistenciesbetweenCUsandex-istingfisherymanagementunits(MUs)(IrvineampFraser2008)withdifferencesbetweenCUsandMUschallengingmanagerswhoareresponsibleforbothassessingbiologicalstatusofCUsandfisheryobjectivesforMUs(HoltampIrvine2013)Giventhelimiteddistribu-tionofcohosalmonpopulationsmarkedwithCWTsinBCitisclearthatCWTscannotprovidetheCUresolutionrecommendedbyPriceetal(2014)Priceetal(2017)forwildpopulationassessmentnortheMUresolutionnotedbyHoltandIrvine(2013)forhatcheryandfisheryassessmentandmanagement
ThecurrentstudyisanevaluationoftheapplicationofthePBT-GSImethodologyoutlinedbyBeachametal(2017)tocohosalmonfisheriesinBCtodeterminewhetherthegenetictechnologiescanbe used to providemore information on fishery contributions byhatcheryandCUthanisavailablefromCWTsCommercialandrec-reationalcohosalmonfisheriesandriverescapementsforselectedpopulationsweresampledforbothCWTsandgenotypesWeeval-uatedthepopulation-levelresolutionobtainedfromCWTsandthegeneticmethodologybyCU for all 2017and some2016 fisheriesinwhichcohosalmonwerecaughtcatchestimationbyCUforthefisheriessampledandstock-specificexploitationrate forselectedpopulationsofcohosalmoninBCGenotypingbysequencingmeth-odologywasusedtogenotypecohosalmonat304singlenucleotidepolymorphisms(SNPs)in304ampliconsCompletebroodstockge-notypingforPBTanalysiswasconductedin2014for20hatchery-enhanced populations that included genotyping 6061 individuals(964genotypingsuccessrate)andastockidentificationbaselinecomprisingsome267populationsrangingfromsoutheastAlaskatoOregonwasemployedforGSIAcomparisonofthepopulation-spe-cificcontributionstomixed-stockfisheriescatchandexploitationratesestimatedwithCWTsandPBT-GSItechnologieswasmadeWeconcludethatageneticapproachcanemulateandimproveupontheresultsavailablefromthecurrentCWTprogramforassessmentandmanagementofcohosalmonenhancementandfisheriesinBCandprovidecriticalinformationtoimprovewildcohosalmonassessmentandconservation
4emsp |emsp emspensp BEACHAM Et Al
2emsp |emspMETHODS
21emsp|emspFishery sample collection
Atotalof8006 individualsweregenotyped fromfisherysamplescollected in2016and2017 In2016sampleswerecollectedfromadiposefin-clippedandunclippedcohosalmonlandedintherecrea-tionalfisheryintheStraitofGeorgiaJuandeFucaStraitandalongthewestcoastofVancouverIslandTheonlyexceptionswereArea21andArea121whereonlyclippedindividualsweresampledandAreas20-1to20-4inAugustwhereonlyclippedindividualsweresampled Sampleswerepooled for analysis asoutlined inTable1withthelocationsofstatisticalareasoutlinedbyhttpswwwpacdfo-mpogccafm-gpmaps-cartesareas-secteursindex-enghtmlAdditionalcohosalmonsamples(bothclippedandunclipped)wereobtained froma troll test fisheryoperatingnearBrooksPeninsulaoff thenorthwest coastofVancouver Island a gillnet test fisherytargetingsockeyesalmon(Oncorhynchus nerka)nearRoundIslandinnorthernJohnstoneStraitandacommercialdemonstrationfreezertrollfisheryinthecentralcoastofBC(Figure1)
In2017forthenorthern(AreaF)freezertrollfisheryselectedfreezerboats(28offleet)wererequiredtokeepheadsofallcohosalmoncaughtwiththemarktype(adiposefinclippedornot)un-knownforanindividualheadUponlandingtheheadswerecountedand checked electronically for CWTs and randomly sampled to amaximumof50headsperdeliveryIfaCWTwasdetectedtheheadwassenttoacentralCWTheadrecoverylaboratoryinVancouverBCwheretheDNAsamplewassubsequentlytakenIfaCWTwassuccessfully recoveredanddecoded froman individualhead from
commercialrecreationalorFirstNationsfisheriesandagenotypewassuccessfullyobtainedforthe individualsample thenthegen-otypesofallof these individualswerepooled intoasinglemixed-stocksampleofknownoriginandknownage inorder toevaluateaccuracyof stock compositionsbyCUandpopulationFieldDNAsamplesweretakenonlyfromindividualswithnoCWTsdetectedInthenortherniceboattrollfisherytheclipstatusofthefishinthecatchwasknownSamplesof clippedcohosalmonwereobtainedfrom this fishery as an ancillary aspect of standard Fisheries andOceansCanadacontractcatchsamplingforCWTsOnlyclippedin-dividualswereexaminedthroughthisprogramwithsimilarsamplingprotocolsasoutlinedinthefreezerboatsamplingInparticularonlyclippedfishnotcontainingaCWTweresampledinthefieldthroughthisprogramwithheadscontainingaCWTsenttotheheadrecov-ery laboratory inVancouverClipped fish constituted16of thecatchinthefisherylargelyduetothewildoriginofthecatchsoitwasimportanttoobtainsamplesfromunclippedindividualsaswellSamplesofunclipped individualswereobtainedby sampling land-ingsforamaximumof25individualspervesselwithamaximumof100 individuals sampled per day Samples from theCentralCoastfreezertrolldemonstrationfisherywereobtainedviaasimilarpro-tocolasoutlinedforthenorthernfreezertrollsamplingTheoriginofthesamplesfromtherecreationalfisheryinBCincludedvolun-taryheadrecoveriesofadiposefin-clippedcohosalmonfromrecre-ationalfisheriesinBCaswellassomecreelsamplingofrecreationalcatches Samples from the recreational fisherywerederived fromclippedindividualsbuttheymaynothavebeenmarkedwithaCWTwhendeliveredtotheCWTheadrecoverylaboratoryThussamplesin2017wereobtainedfromall individualsthatwouldberoutinely
Fishery Reporting region Statistical area
Recreational JohnstoneStrait(JST) A11toA13
StraitofGeorgia-north(SOG-n) A14toA16
StraitofGeorgia-south(SOG-s) A17toA18A19-7toA19-12A28A29
StraitofGeorgiaJuandeFucaStrait(SOGJDF)
A19-1toA19-6A20-5
WestcoastVancouverIslandJuandeFucaStrait(WCVIJDF)
A20(exceptA20-5)
SouthwestVancouverIsland(SWVI21121)
A21A121
SouthwestVancouverIsland-inshore(SWVI-inshore)
A23A24
SouthwestVancouverIsland-offshore(SWVI-offshore)
A123A124
NorthwestVancouverIsland-inshore(NWVI-inshore)
A25toA27
NorthwestVancouverIsland-offshore(NWVI-offshore)
A125toA127
Test BrooksPeninsula A27A127A126
RoundIsland A12
Troll CentralCoast A6A7A8
TA B L E 1 emspReportingregionandDFOstatisticalareaswithinregionsforamalgamationofcatchsamplesLocationsofstatisticalareasareoutlinedonhttpswwwpacdfo-mpogccafm-gpmaps-cartesareas-secteursindex-enghtml
emspensp emsp | emsp5BEACHAM Et Al
processed under the CWT recovery program for coho salmon inBCInadditionsampleswerecollectedfromarecreationalfishery(clippedandunclipped individuals) fromHaidaGwaiinearLangaraIslandarecreational fisherynearRivers Inlet (Area8clippedandunclippedindividuals)inthecentralcoastthepreviouslynotediceboat northern troll fishery and direct creel sampling (clipped andunclippedindividuals)ofrecreationalfisheriesinsouthernBC
Monthly catch composition estimates were determined forclipped individuals from various fisheries in BC Seasonal catchcomposition estimateswere determined by recording the numberofindividualgenotypesavailablefromthefisherysamplingineachmonth andweighting individualmonthly samples by the recordedcatch in themonth Individualswere randomlychosen formonthsinwhich the number of available genotypeswas in excess of thenumberrequiredfortheseasonalsampleForexampleif60oftheannualcatchwasrecordedinonemonthand200genotypeswereavailablefromthefisherythenallindividualssampledwouldbein-cludedintheseasonalsamplebutcontributionsofindividualsfromothermonthswerescaledrelativetothemonthlycontributiontothe
seasonalcatchIfthecatchinoneothermonthwas20oftheannualcatchand150sampleswereavailablethen200(2060)=67genotypeswererandomlychosenfromtheavailable150genotypesforinclusionintotheseasonalsample
InthenorthernBCtrollfisherysamplerselectronicallycheckedclippedindividualsforthepresenceofaCWTandheadsfromthoseindividualscontainingaCWTweresubsequentlysenttothecentrallaboratory for CWT recovery and tissue sampling for subsequentgenotypingTissuesamplesfromclippedindividualswithnoCWTsweredirectlyprovidedforgenotypingInordertoestimatemonthlyandseasonalstockcompositionofclippedindividualsinthisfisherywepooledthegenotypesobtainedfromdirectfisherysamplingofclippedindividualswiththosesubsequentlysenttothecentrallabo-ratoryforCWTrecoveryinordertoprovideanappropriatepoolofclippedindividualstoestimatemonthlyandseasonalstockcompo-sitionsofclippedindividualsinthefisheryIneachsampleanalyzedover all BC fisheries thenumberof individuals identified viaPBTrelative to thenumberofgenotypes in the samplewas tabulatedandsummarizedoversamplefisheryandseason
F I G U R E 1 emspMapindicatinggeographiclocationsforfisherysamplingand20populationsforwhichparentage-basedtaggingwasappliedinestimationofstockcomposition
6emsp |emsp emspensp BEACHAM Et Al
22emsp|emspEscapement sample collection
In2016adiposefin-clippedjacks(age2years)weresampledfromeightpopulationswith573individualscollectedandgenotypedsuc-cessfullyJacksweredefinedonthebasisofbodysizeduringsam-plecollectionwith jacksvisiblysmallerthanage3-yearspawnersPopulations sampled were as follows Robertson Creek QuinsamRiver Puntledge River Big Qualicum River Goldstream RiverChilliwackRiverStaveRiverand InchCreekTheobjectiveofthesamplingwas to obtain a sample of known origin and known agebaseduponknowncollectionlocationinordertoevaluateaccuracyofassignmentsof the jacks tobothCUandpopulationunder theassumption of no straying among populations for the individualssampled
In 2017 escapements (non-broodstock hatchery and river re-turns clipped individualsonly)were sampledwhere samplingwasfeasibleEscapementsamplingfrom13populationswasconductedand1692 individualsweregenotypedEscapement sampleswerenot available from all 20 populations forwhich itwas possible toassignindividualsinthenon-broodstockescapementsviaPBTTheobjectiveofthesamplingwastoevaluatestrayingratesamongpop-ulationsandforthosehatcheries(ChilliwackRiver)whereparentalgenotypeswereassignedtospecificreleasegroupsprovide infor-mationon the relative rates of returnof different release groupsAs relative ratesof return for the releasegroupswereconsideredancillarytothemainfocusofthestudynoresultsfromthesereturnswereincludedinthecurrentstudy
23emsp|emspExploitation rate
WeestimatedexploitationrateofadultcohosalmoninBCfisheriesviabothCWTsandgeneticsExploitationrateforapopulationwasdefinedasadultcatch(adultcatch-escapement)ForCWTstheob-servednumberofCWTswascorrectedbyldquono-pinrdquotaglossratesandwasexpandedbythepopulationrsquostag-specificmarkingratesummedovertagcodesandexpandedagainbythesamplingrateforthefish-eryinordertoestimatecatchofhatchery-originindividualsTheob-servednumberofCWTsinescapementsamplingwasexpandedinasimilarmanner inordertoestimatethehatcherycontributiontothe escapement For genetics the seasonal kept catchof adiposefin-clippedindividualsinafisherywasmultipliedbyaseasonalstockcompositionestimateinordertoestimatepopulation-specificcatchTheabundanceofhatchery-originescapementwascalculatedastheestimatedescapementmultipliedbytheproportionofadiposefin-clippedadultsobserved in theescapement If partof the juvenileproductionwasnotadiposefinclippeduponreleasefromthehatch-erythenthehatcherycontributionstobothcatchandescapementwereunderestimatedbythismethod
24emsp|emspGenetic stock identification baseline
The initial baseline was outlined by Beacham et al (2017) andconsisted of 20242 individuals from 117 populations with the
distributionofpopulationsrangingfromsoutheastAlaskatoPugetSoundinWashingtonStateThebaselinehasbeensubsequentlyex-panded to include 40774 individuals from267populations rang-ingfromsoutheastAlaskatoOregonTheprimaryexpansionofthebaselineincludedasurveyofadditionalpopulationsinsouthernBCcoastalWashingtontheColumbiaRiverdrainageandOregonThefull baseline is outlined in Supporting Information Table S1 withthe populations fromBC arrangedbyCU andwithUnited StatesofAmerica(US)populationsarrangedbygeographic(reporting)re-gionBeachametal (2017)hadpreviouslydemonstratedastrongregionalpopulationstructureandreportedthatoverall117popula-tionsaverageself-assignmentaccuracytopopulation(weightedbypopulation sample size)was 855 for individualswith an assign-mentprobabilitylt050excludedand928forindividualswithanassignmentprobabilitylt085excludedHighlevelsofaccuracyforGSI assignment of individuals to specific populations when com-binedwithassignmentsviaPBTprovideapowerful technique forassignmentofindividualsofunknownorigintosomepopulations
25emsp|emspLibrary preparation and genotyping
ThedetailedprocedureforlibrarypreparationandgenotypingwasoutlinedbyBeachametal(2017)SummarizedbrieflyDNAwasex-tractedfromcohosalmontissuesamplesviaaChelexextractionandtheDNAconcentration normalized to40ngμlwith a Tecan LiHarobotApanelofprimersdesignedtoamplifyspecificsegmentsofDNAthatcontainedSNPsofinterestwasappliedtotheextractedDNAwithprimersequencesforeachampliconoutlinedbyBeachametal (2017SupportingInformationTableS2)TheinitialmultiplexPCR amplification of 304 target ampliconswas conductedwith acocktail of 2μl of normalizedDNA extract 5μl of 2X Ion Agriseqprimerpool2μlofIonAgriseqHiFimixand1μlofddH2O Thermal cyclingwasconductedin96-wellPCRplates(oneindividualperwell)withthefollowingconditionsforPCR99degCndash2min17cycles(99degCndash15s60degCndash4min)10degChold
FollowingtheinitialPCRasecondstepemployingathermalcy-clerwasconducted thatpartiallydigested theprimerson theam-pliconsandthereactionconductedwiththefollowingconditions50degCndash10min55degCndash10min60degCndash20min10degCholdA thirdandfinalstepemployingathermalcyclerwasinitiatedtoligatethebarcodes(768individualcodes)totheampliconsandwasconductedwiththefollowingconditions22degCndash30min70degCndash10min10degCholdLibrarieswerepurifiedbyadditionof225μlofAgencourtregAMPureregXPmagneticbeadstoeachlibrarytheplatewasplacedonamagnetic racksupernatantdiscardedandthebeadswashedtwice in70ethanolThepurified librarieswerethenelutedwith25 μloflowTEand20μlofthesupernatanttransferredtoafresh96-well trayNext eachof the768prepared librarieswaspooledintoasingletubeforprocessingontheIonChefreg(ThermoFisherScientific)Twotubesofpooled librarieswereprocessedconsecu-tivelyontheIonChefandthus1536individualswereprocessedon a single run of the Ion ChefOne tube of the pooled librarieswasloadedontoeachP1regchipv3andthusampliconsfrom768
emspensp emsp | emsp7BEACHAM Et Al
individualsweredistributedoneachP1chipwith1536individualsprocessedbetweentwochipsThechipswerethenloadedontotheIonTorrentProtonsequencerAfter thesequencingrunwascom-pletedcomparisonswiththereferencegenomeoftherainbowtrout(O mykiss)(Berthelotetal2014)supplementedwiththesequencescontaining theobservedcohosalmonSNPswereconductedwithProton softwareVariant Callerreg and SNP genotypes at the sitesspecified by the hotspot filewithin target regionswere called byVariantCallerThehotspotfilecontained304SNPsiteswithoneSNPscoredateachampliconGenotypesatallavailableSNPsforanindividualwereassembled toprovideamulti-locus individualgen-otype The species identification SNPOkiOts_120255‐113 (StarksClementoampGarza2016)andsex identificationSNPOts_SEXY3‐1 wereomittedfromsubsequentparentageandGSIanalysesleaving302SNPs
26emsp|emspIdentification of individuals
AsnotedpreviouslyPBTandGSIwereusedconcurrently toesti-mate fishery stock composition Initially PBT was used and theanalysiswas conductedwhere the genotypes of individuals to beidentifiedwerematched to the genotypes of prospective parents(COLONY JonesampWang 2010Wang 2016) If all individuals ina hatchery broodstock are sampled and subsequently genotypedthenalloffspringfromthebroodstockaregeneticallymarkedThegenotypes of individuals of unknown origin are statistically com-pared with the genotypes of potential parents and if a match ismadetheoffspringareassignedtotheparentsandthustheori-ginandageoftheindividualsaredeterminedParentageassignmentsoftwarewasutilizedtoassignoffspringtoparentsasCOLONYcanproduceassignmentswhen thegenotypeofoneof theparents ismissingeitherduetoamissingparentalsampleor failure topro-duceaparentalgenotypefromanexistingsampleGiventhatPBTassignments for20potential populationswereevaluated for eachfisherysampleCOLONYwasrunwithallbroodstocksampleddur-ing2014inputasasingleunitforanalysisoffisherysampleswithnodifferentiationamongpopulationsAlthoughtheCOLONYassump-tionofasinglepopulationintheparentpoolwasviolatedanalysisofknown-originsamplesindicatedthatveryhighlevelsofaccuracywereachievedinassignmentswhenpoolingofpotentialparentsincontributing populationswas conducted Two-parent assignmentswere accepted only when both assigned parents originated fromthesamepopulationotherwisetheindividualwaspassedtogeneticstockidentification(GSI)forpotentialassignmentTwo-parentandsingle-parentassignmentswereacceptedonlywhentheprobabilityofcorrectassignmentwasge085fortheparentpairotherwisetheindividualwaspassedtopotentialassignmentbyGSIAdditionallyforsingle-parentassignments tobeaccepted thePBTassignmenttopopulationhad tobepart of theCUassigned to the individualviaGSI Individuals forwhichnoprospectiveparentswere identi-fiedinthebroodstockwerepassedtoGSIforpotentialassignmentPolygamousmatingwasassumedfortheCOLONYanalysisSimplepairwisecomparisonsbetweenoffspringandpotentialparentswere
conductedThebaselineforindividualssampledinthe2016escape-ments (jacks)and2017fisheries includedallbroodstockssampledin2014as these individualsarepredominately threeyearsofage(Sandercock1991)Jacks inthe2017escapementwere identifiedvia body size and subsequent assignment to parents in the 2015hatcherybroodstocksIndividualswithmorethan120missinggeno-typeswereeliminatedfromfurtheranalysesAnestimatedgenotyp-ingerrorrateof1wasusedforCOLONYassignmentsPreviouslyBeachametal(2017)hadreportedthatanaveragegenotypingerrorrate of 107 (1220discrepancies in 114105 comparisons) or analleleerrorrateof053(1220discrepanciesin228210compari-sons)wasobservedoverthe302SNPsscoredTheparentpairout-putfilewasthebasicfileusedinsubsequentanalyses
Thesecondmethodof individual identification isGSI inwhichthegeneticprofilesofwholepopulationspotentiallycontributingtoamixed-stocksampleareusedtoestimatetheoriginofeachindivid-ualinthesample(RUBIASMoranampAnderson2018)ThisanalysiswasrestrictedtothoseindividualsunassignedviaCOLONYForeachsample individuals not assigned by COLONYwere then assignedwithRUBIASThepopulationposteriormeansfilewasthebasicfileusedforsubsequentanalysesandthisfilecontainedtheprobabilityofassignmentoftheindividualtoeachofthe267populationsinthebaselineStockcompositionwasestimatedthroughthecombinationoffilesgeneratedwithbothCOLONYandRUBIAS Individualsas-signedviaCOLONYwereassignedaprobabilityof100oforiginat-ingfromtheidentifiedpopulationwitha000probabilityassignedtoallotherpopulationsinthebaselineThislevelofassignmentac-curacyviaPBTwasobservedpreviouslyforcohosalmonwiththepanelofSNPsemployedinthecurrentstudy(Beachametal2017)ThesedatawerethencombinedwiththeprobabilityofassignmentforthoseindividualsunassignedviaCOLONYtoeachofthepopu-lationsinthebaselineviaRUBIASAfteraninitialburn-inof25000iterations the last1000 iterations from theMonteCarloMarkovchainfromRUBIASwereusedtoestimatetheoriginofindividualsandstockcompositionwiththemeanallocationtoeachpopulationinthebaselinedeterminedStandarddeviationsofestimatedstockcompositionswere alsodetermined from the last1000 iterationsfrom theMonteCarloMarkovchain Stock compositionbyCUorreportinggroupwasdeterminedbysummationofallocationstoallpopulations in the baseline that belonged to the CU or reportinggroupunderconsideration
3emsp |emspRESULTS
31emsp|emspEstimation of stock composition for known‐origin samples
Genotypeswere available from573 jacks sampled in2016acrossthreeCUs and eight populations in southernBC Estimated stockcompositionbyCUand reportinggroup for this combinedsamplewas accuratewith an error utilizingonlyGSI of le04byCU forthreeCUspresentinthesample(Table2)WithbothPBTandGSIutilized theaverageerrordeclinedtole02byCUfor threeCUs
8emsp |emsp emspensp BEACHAM Et Al
TAB
LE 2
emspAccuracyofregional(UnitedStates)andconservationunit(Canada)estimatedstockcomposition()foramixed-stocksampleof573known-origincohosalmonjackssampledin
2016and620coded-wiretaggedindividualssampledin2017fisheriesinBCStandarddeviationoftheestimateisinparentheses
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
SoutheastAlaska
00
00(00)
00(00)
90
68(11)
68(11)
AlsekRiver
00
00(00)
00(00)
00
00(00)
00(00)
LowerStikine
00
00(00)
00(00)
00
04(03)
04(03)
LowerNass
00
00(00)
00(00)
63
64(10)
62(10)
UpperNass
00
00(00)
00(00)
00
04(03)
04(04)
PortlandSound-Observatory
Inlet-PortlandCanal
00
00(00)
00(00)
00
00(00)
00(00)
SkeenaEstuary
00
00(00)
00(00)
00
00(00)
00(00)
LowerSkeena
00
00(00)
00(00)
16
12(05)
12(05)
MiddleSkeena
00
00(00)
00(00)
69
39(08)
39(09)
UpperSkeena
00
00(00)
00(00)
06
40(10)
40(09)
HaidaGwaiindashGrahamIslandLowlands
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-East
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-W
est
00
00(00)
00(00)
00
00(00)
00(00)
NorthernCoastalStreams
00
00(00)
00(00)
00
00(00)
00(01)
HecateStraitMainland
00
00(00)
00(00)
00
07(05)
06(04)
MusselndashKynoch
00
00(00)
00(00)
00
00(00)
00(00)
DouglasChannelndashKitimatArm
00
00(01)
00(01)
00
00(01)
00(01)
BellaCoolandashDeanRivers
00
00(00)
00(00)
00
00(00)
00(00)
RiversInlet
00
00(01)
00(00)
00
00(00)
00(00)
SmithInlet
00
00(00)
00(00)
00
00(00)
00(00)
SouthernCoastalStreamsndashQueen
CharlotteStraitndashJohnstoneStraitndash
SouthernFjords
00
00(01)
00(01)
00
00(02)
00(02)
HomathkondashKlinakliniRivers
00
00(00)
00(00)
00
00(00)
00(00)
GeorgiaStraitMainland
00
00(00)
00(00)
00
00(00)
00(00)
HoweSoundndashBurrardInlet
00
00(00)
00(00)
06
00(01)
00(00)
EastVancouverIslandndashGeorgiaStrait
351
345(20)
348(20)
131
134(14)
140(14)
EastVancouverIslandndashJohnstone
StraitndashSouthernFjords
00
00(00)
00(00)
00
00(01)
00(00)
NahwittiLowland
00
00(01)
00(00)
18
17(06)
17(06) (C
ontin
ues)
emspensp emsp | emsp9BEACHAM Et Al
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
WestVancouverIsland
115
115(14)
115(13)
135
134(14)
135(14)
Clayoquot
00
00(00)
00(00)
00
02(03)
02(02)
JuandeFucandashPachena
00
00(01)
00(01)
00
00(00)
00(00)
LowerFraser
534
539(21)
536(21)
185
187(15)
188(15)
Lillooet
00
00(00)
00(00)
00
03(02)
02(01)
FraserCanyon
00
00(00)
00(00)
00
00(00)
00(00)
InteriorFraser
00
00(00)
00(00)
00
00(00)
00(00)
LowerThompson
00
00(00)
00(00)
08
07(03)
08(04)
NorthThompson
00
00(00)
00(01)
00
00(00)
00(00)
SouthThompson
00
00(00)
00(00)
05
06(03)
05(03)
Boun
dary
Bay
00
00(00)
00(00)
00
00(00)
00(00)
NooksackRiver
00
01(02)
01(02)
26
09(09)
10(08)
SkagitRiver
00
00(01)
00(01)
05
23(08)
22(08)
NorthernPugetSound
00
00(00)
00(00)
39
58(12)
55(13)
Mid-PugetSound
00
00(00)
00(00)
52
62(11)
63(11)
SouthernPugetSound
00
00(00)
00(00)
10
09(10)
06(08)
JuandeFucaStrait
00
00(00)
00(00)
00
02(03)
01(03)
HoodCanal
00
00(00)
00(00)
44
16(05)
16(06)
CoastalWashington
00
00(00)
00(00)
60
59(10)
59(10)
ColumbiaRiver
00
00(00)
00(00)
32
32(07)
32(07)
Oregon
00
00(00)
00(00)
00
00(01)
00(00)
TAB
LE 2
(Continued)
10emsp |emsp emspensp BEACHAM Et Al
presentinthesample(Table2)Theaverageerrorfor45CUsorre-portinggroupsabsentinthesamplewas00bybothGSIandPBT-GSIBypopulation543(948)ofthejackswereassignedviaPBTwith 100 accuracywith respect to population of origin and age(Table3)Estimatedstockcompositionfor20populationswhereitwaspossibletousebothPBTandGSIinestimationofstockcompo-sitionwasaccurateFortheeightpopulationspresentinthesampleofjackstheaverageerrorutilizingGSIonlywas05perpopulationdecliningto02utilizingbothPBTandGSIundertheassumptionthatthecollectionlocationwasanaccuratereflectionofjackoriginTherewasanaverage00errorforthe12populationswithnorep-resentation inthesamplewithutilizationofeitherGSIorPBT-GSI(Table4)ThelargesterrorutilizingbothPBTandGSI(underestima-tionby06ofactualvalue)wasobservedintheGoldstreamRiverpopulationwhichalsodisplayedthe lowestpercentage (704)ofjacksidentifiedviaPBT(Table3)PBTwhencombinedwithGSIpro-ducedmoreaccurateestimatesofCU-specificandpopulation-spe-cificstockcompositionthanavailablewithonlyGSI
Of the 3054 adipose fin-clipped coho salmon samples re-ceivedforpotentialgenotypingfromacentrallaboratorywhere2017fisherysampleswereprocessedforpotentialCWTrecov-erygenotypeswereobtained for2533 (829)of the individ-uals processed There were 750 CWTs recovered from 3054headsor snouts examined and genotypeswereobtained from622 (829) of the individuals that carried a CWT GenotypesofcohosalmonwithoutCWTsbutsentfromtheCWTheadre-covery laboratorywereobtainedfrom1911of2304 individu-als (829) Release information associatedwith theCWTwasavailable for 620 of the 622CWTs associatedwith genotypedindividuals and these 620 individuals constituted a sample ofknown origin There were individuals from 20 regions or CUspresentinthissampleandforthese20regionsorCUstheav-erageerrorofestimationutilizingonlyGSIwas09perregionorCUand09forbothPBTandGSIcombined(Table2)CWTswererecoveredoriginatingfromsevenpopulationswhereitwaspossibletoincorporatebothGSIandPBTinestimationofstockcompositionFor these sevenpopulations theaverageerror inestimatedstockcompositionsperpopulationwas20withGSIand 03 with PBT-GSI (Table 4) There was an average 00errorforthe13populationswithnorepresentationinthesam-ple with utilization of either GSI or PBT-GSI (Table 4) Similartothesampleof jacksPBTwhencombinedwithGSIproducedmore accurate estimates of population-specific stock composi-tionthanavailablewithonlyGSI
32emsp|emspEvaluation of 2016 fishery sampling
Asbroodstocksamplingatselectedhatcheries insouthernBCdidnot commence until 2014 it was not possible to obtain PBT as-signments for any fishery samples collected in 2016 HoweverGSI analysis of individuals derived from recreational fishery sam-ples intheStraitofGeorgiaJuandeFucaStraitandoffthewestcoastofVancouverIslandinsouthernBCindicatedthattherewas
virtually no contribution from regions orCUsnorthofVancouverIsland (Supporting Information Table S2) The only exceptionwasin thenorthwestcoastofVancouver Islandboth inshoreandoff-shorewhereupto12ofthesamplewasidentifiedasoriginatingfromnorthernandcentralBCCUsUS-origincohosalmonwerees-timatedtohavecontributedapproximately70ofsummerrecrea-tionalsamplesderivedfromoffshorefisheriesalongthewestcoastofVancouverIslandaswellasintheentrancetoJuandeFucaStrait(A21A121)InmoreinshoreareasalongthewestcoastofVancouverIslandcohosalmonfromlocalCUswerethedominantcontributorstothefisheryForexamplethewestcoastofVancouverIslandCUcontributedapproximately80ofthe39-individualAugustsamplefromtheinshoresouthwesternVancouverIslandregionIntheStraitofGeorgia localCUswereagainthemaincontributorstotherec-reationalfisheryduringthesummerwiththeHoweSoundndashBurrardInletCUcontributingupto40insomesamplestheeastcoastofVancouverIslandCUuptoapproximately30andthelowerFraserRiverCUuptoapproximately40WhilethelowerFraserRiverCUwasanimportantcontributortotherecreationalfisheryintheStraitofGeorgiaandJuandeFucaStraititscontributiontofisheriesoffthewestcoastofVancouverIslandwaslimitedTheNooksackRiverregion innorthernWashingtonStatewasalso identifiedasacon-sistent contributor to recreational fishery samples in the Strait ofGeorgia insomecasescontributingupto20ofthesampleana-lyzed(SupportingInformationTableS2)
StockcompositionswereavailablefromtwotestfisheriesoneatrolltestfisherynearBrooksPeninsulabynorthwesternVancouverIslandandtheotheragillnettestfisheryinupperJohnstoneStraitIn the Brooks Peninsula fishery Vancouver IslandCUs comprised40ndash60ofthesampleswithaminorcontributionofFraserRiverCUs (lt10) The five interior Fraser River CUs of conservationconcern comprised00ndash33of the sampleswhereas themoreabundant lower Fraser River CUs comprised only 00ndash55 ofthesamples Individuals fromFraserRiverCUswerenotdetectedduringJulyinArea127theoffshoreportionofnorthernnorthwest-ernVancouverIslandbuttheyweredetectedinArea126themoresoutherlyoffshoreportionofnorthwesternVancouverIslandIntheJohnstoneStrait(RoundIsland)testfisheryindividualsfromFraserRiverCUswerescarcebutthefiveinteriorFraserRiverCUscom-prised30ndash35of the sampleswhereas the lowerFraserRiverCUscomprisedonly00ndash65ofthesamplesThemajorcontrib-utors to the samplesoriginated from theeast coastofVancouverIsland(24ndash45)andHoweSoundndashBurrardInletCUs(9ndash11)aswellascoastalWashington(8ndash15)(SupportingInformationTableS2)
ThefinalfisherysampledwasademonstrationcommercialtrollfisheryincentralcoastalBC(A6andA7)ContributionsfromlocalCUswerehigherinsamplesfrommoreinshoreareas(A7-7August212southernCUsandregions)withcontributionsfromsouth-ernCUshigherinthemoreseawardareas(A6-9September862southernCUsandregions)LikethewestcoastofVancouverIslandmigrating stocks were more likely to be found in more seawardlocations
emspensp emsp | emsp11BEACHAM Et Al
33emsp|emspEvaluation of 2017 fishery sampling
Onesourceoffisherysampleswasdirectfisherysamplingofindi-viduals(bothadiposefin-clippedandunclipped)wherethefocuswas not associated with potential CWT recovery In this casesamplingofunclippedindividualsfromthenorthern(AreaF)trollfishery indicated that initially (July 10ndash12) northern and centralBCCUscontributed23tothecatchsouthernBCCUs53andsouthern US regions 23 (Supporting Information Table S3)Stock composition changed markedly in samples collected July13ndash18 with Alaska contributing 12 northern and central BCCUs contributing 72 and southern BCCUs contributing 16withnosouthernUScontributionSamplescollectedduringJuly22ndash28revealedadominantcontributionfromnorthernandcen-tralBCCUs(96)andCUsfromthisregioncontributedbyfarthelargestportionof the catch in subsequent sampleswith south-ernBCCUsandsouthernUSregionslargelyabsent(SupportingInformation Table S3) The average contribution to the weeklysamples from July 22 onwardswas 17 southeast Alaska 16HaidaGwaiindashGrahamIslandLowlandsCU15DouglasChannelndashKitimatArmCU 12Hecate StraitMainlandCU 8NorthernCoastalStreamsCUand7LowerNassCUNotunexpectedlysampling of only clipped individuals from the fishery indicatedverysubstantialdifferencesinstockcompositionbetweenclippedandunclippedindividualsTheaveragecontributiontotheweeklysamples of clipped individuals from July 19 onwards was 32coastalWashington11mid-PugetSound10westVancouverIsland CU and 5 for each of northern Puget Sound and Juande Fuca Strait (Supporting Information Table S4) However theobservedcliprateinthefisherywas16sothecatchwasdomi-natedbyunclippedindividualsTheSeptember2ndash8samplefromthe Haida Gwaii recreational fishery near Langara displayed astockcompositionof42HaidaGwaiindashGrahamIslandLowlandsCU15HecateStraitMainlandCU10LowerSkeenaCUand8 Northern Coastal Streams CU similar to that observed inthe troll fisherywith the local CU (HaidaGwaiindashGraham IslandLowlands) inhigherproportionsaswouldbeexpectedinamoreinshorefishery(SupportingInformationTableS3)
Directsamplingof thecentralcoast troll fisheryrevealedthatonaverage theDouglasChannelndashKitimatArmCUwasthe largestcontributortothefishery(24)followedbytheNorthernCoastalStreamsCU(19)BellaCoolandashDeanRivers(18)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(10) (Supporting InformationTableS3)withallCUsproximaltothelocationofthefisherySamplesfromtherecreationalcentralcoastfisheryindicatedthattheBellaCoolandashDeanRiversCUwasthedominant contributor to the fishery (32) followedby theRiversInletCU(19)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(15)andDouglasChannelndashKitimatArmCU(7) (SupportingInformationTableS3)GiventhelocationofthefisherydominantcontributionsfromtheseCUsweretobeexpected
AsnotedpreviouslysampleswerealsoavailablefromvirtuallyallindividualssampledfromcommercialrecreationalandFirstNationsfisheriesthatwouldberoutinelyprocessedforCWTrecoveryinayearbyacentrallaboratoryIndividualheadsfromcommercialfish-eries sent to the central laboratoryhadbeenpreviously screenedelectronically for thepresenceofaCWTVoluntaryheadsamplesfromadiposefin-clippedindividualsfromrecreationalfisheriesweresenttothecentrallaboratorybutmanyoftheseindividualsdidnotcontainaCWTastherewasnopriorscreeningof these individu-als Stock composition of clipped individuals in the northern trollfishery has been previously outlined above Stock composition inthenorthern recreational fishery indicatedawidegeographicdis-tributionofcontributions rangingfromcoastalWashington (16)thewestcoastofVancouverIslandCU(10)theupperSkeenaCU(9)southeastAlaskaregion(8)mid-PugetSound(7)andeastcoast of Vancouver IslandndashStrait of Georgia CU (6) (SupportingInformation Table S4) Stock composition of clipped individuals inthecentralcoastrecreationalfisherywasdominatedbytheHoweSoundndashBurrardInletCU(27)andeastVancouverIslandndashStraitofGeorgiaCU(25)(SupportingInformationTableS4)butgiventhelowobservedclip rate in the fishery (014) therewasvery littlecatchthatcouldbeidentifiedashatcheryorigin
Directsamplingofrecreationalfisheries insouthernBCwhichincluded both clipped and unclipped individuals in the sample
Population Genotyped Assigned assignment
accuracy of population assignment
Robertson 66 66 1000 1000
Quinsam 69 68 986 1000
Puntledge 21 19 905 1000
BigQualicum 84 84 1000 1000
Goldstream 27 19 704 1000
Chilliwack 195 183 938 1000
Inch 91 85 934 1000
Stave 20 19 950 1000
Total 573 543 948 1000
TA B L E 3 emspPercentageofadiposefin-clippedjacks(age2years)sampledin2016andassignedviaparentageanalysistoparentsin2014broodstockpopulationsPopulationofsamplingwasassumedtobethepopulationoforiginofthejacksforevaluationofaccuracyofassignment
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp3BEACHAM Et Al
was33andthoseofCanadianoriginwas25oftherespectiveannual numbers for the base period largely due to a substantialreductionincatchbutalsocoincidentwithreductioninthenum-berofCWTsappliedSinceestimationoffisheryimpactswiththeFRAM is dependent on CWT recovery information the greatlydecreased number of CWT recoveries in recent years will haveincreased the variance of the estimated stock-specific catch andexploitationratesinfisheries(Hinrichsenetal2016ReisenbichlerampHartmann1980)
Anewcost-effectiveapproachtotheassessmentandmanage-mentofwildcohosalmonandtheassociatedhatcheryproductionand fishery management is needed Anderson and Garza (2006)noted that parentage-based tagging (PBT) provides equivalent in-formation (hatcheryof releaseageof individual) forhatchery fishasdoCWTsimplementationofPBTthusmayovercomeproblemsassociatedwithCWT-basedassessmentandmanagementof cohosalmon fisheries inBCAdditionally PBTprovides ameansof im-provedhatcherybroodstockmanagementaswellasassessmentofhatchery-wild interactions in salmonids Unlike CWT-based man-agement PBT-informed hatchery and fishery management wouldbenefit from the complete adipose-clipping of hatchery-producedsalmonAsignificantadvantageofthecombinationofmassmarkingandPBT implementation is thecapabilityto identifyvisuallysam-ple and if desired remove hatchery fish of local and stray originin threatenedwildpopulationsMoreover PBTentails genotypingtheentirehatcherybroodstockandenablestheidentificationofallhatcheryprogenybyparentageassignment(Anderson2012Wang2016)thusenablingaldquomarkraterdquoofvirtually100ofhatcheryfishSteeleetal(2013)demonstratedtheequivalencyofCWTandPBTinaninitialevaluationofpopulationandageassignmentinsteelheadtrout(Oncorhynchus mykiss)oftheSnakeRiverbasinintheColumbiaRiverdrainageHessetal (2016)expandedtheapproachbyusingboth PBT and genetic stock identification (GSI) to investigate runtiming of steelhead trout in the upper Columbia River drainageTheseapplicationsconfirmedthecapabilityofacombinedPBT-GSItechnologytoprovideequivalentorbetter identificationoffishastheCWTmethodbutwerelimitedingeographicscaleInthisstudyweexaminewhether theexistingPBTandGSI systemof identifi-cationforCanadian-origincohosalmon(Beachametal2017)canprovidetheinformationrequiredforimprovedassessmentandman-agementofcohosalmonmixed-stockfisheriesinBritishColumbiacoveringamuchbroadergeographicrangeofcohopopulationsfromAlaskatoOregon
Although propermanagement of hatchery production and as-sociated fishery exploitation requires properly defined objectivesandmonitoring (Flagg2015) it is not currentlypossible toevalu-ate each Canadian enhancement project separately (TompkinsHamiltonBatemanampIrvine2016)ForcohosalmonCWTsarenotappliedtoreleasesfromsomeofthelargesthatcheriesinsouthernBC(ChilliwackRiverCapilanoRiverChehalisRiverConumaRiverNitinatRiverandTenderfootCreek)duetofundinglimitationsandthus theirspecificcontributionstohighlymixed-stockoceanfish-eriesareunknownInfactCWTsareappliedonlytocohosalmon
juvenilesreleasedfromsixFisheriesandOceansCanadalargepro-duction hatcheries three on the east coast of Vancouver Island(QuinsamRiverPuntledgeRiverandBigQualicumRiver)oneonthewestcoastofVancouverIsland(RobertsonCreek)oneinthelowerFraserRiverdrainage(InchCreek)andoneintheThompsonRiver(amajortributaryoftheFraserRiver)drainage(SpiusCreekwheretheSalmonRiverandEagleRiver juvenilesmaybe reared)CWTsarealsoappliedatsmallerfacilities(SeymourRivernearVancouverTobogganCreekandZymacordRiver intheSkeenaRiverdrainageinnorthernBC) and somenaturally spawned indexpopulations innorthernBC
The 43 coho salmon CUs originally identified by Holtby andCiruna(2007)havebeenmodifiedtoacurrentnumberof44CUsPrice Rosenberger Taylor and Stanford (2014) Price EnglishRosenbergerMacDuffeeandReynolds (2017)suggestedthatanysuitable assessment technique must provide individual resolutionfor all CUs to meet the conservation requirements of CanadarsquosWSPTherecanbepossible inconsistenciesbetweenCUsandex-istingfisherymanagementunits(MUs)(IrvineampFraser2008)withdifferencesbetweenCUsandMUschallengingmanagerswhoareresponsibleforbothassessingbiologicalstatusofCUsandfisheryobjectivesforMUs(HoltampIrvine2013)Giventhelimiteddistribu-tionofcohosalmonpopulationsmarkedwithCWTsinBCitisclearthatCWTscannotprovidetheCUresolutionrecommendedbyPriceetal(2014)Priceetal(2017)forwildpopulationassessmentnortheMUresolutionnotedbyHoltandIrvine(2013)forhatcheryandfisheryassessmentandmanagement
ThecurrentstudyisanevaluationoftheapplicationofthePBT-GSImethodologyoutlinedbyBeachametal(2017)tocohosalmonfisheriesinBCtodeterminewhetherthegenetictechnologiescanbe used to providemore information on fishery contributions byhatcheryandCUthanisavailablefromCWTsCommercialandrec-reationalcohosalmonfisheriesandriverescapementsforselectedpopulationsweresampledforbothCWTsandgenotypesWeeval-uatedthepopulation-levelresolutionobtainedfromCWTsandthegeneticmethodologybyCU for all 2017and some2016 fisheriesinwhichcohosalmonwerecaughtcatchestimationbyCUforthefisheriessampledandstock-specificexploitationrate forselectedpopulationsofcohosalmoninBCGenotypingbysequencingmeth-odologywasusedtogenotypecohosalmonat304singlenucleotidepolymorphisms(SNPs)in304ampliconsCompletebroodstockge-notypingforPBTanalysiswasconductedin2014for20hatchery-enhanced populations that included genotyping 6061 individuals(964genotypingsuccessrate)andastockidentificationbaselinecomprisingsome267populationsrangingfromsoutheastAlaskatoOregonwasemployedforGSIAcomparisonofthepopulation-spe-cificcontributionstomixed-stockfisheriescatchandexploitationratesestimatedwithCWTsandPBT-GSItechnologieswasmadeWeconcludethatageneticapproachcanemulateandimproveupontheresultsavailablefromthecurrentCWTprogramforassessmentandmanagementofcohosalmonenhancementandfisheriesinBCandprovidecriticalinformationtoimprovewildcohosalmonassessmentandconservation
4emsp |emsp emspensp BEACHAM Et Al
2emsp |emspMETHODS
21emsp|emspFishery sample collection
Atotalof8006 individualsweregenotyped fromfisherysamplescollected in2016and2017 In2016sampleswerecollectedfromadiposefin-clippedandunclippedcohosalmonlandedintherecrea-tionalfisheryintheStraitofGeorgiaJuandeFucaStraitandalongthewestcoastofVancouverIslandTheonlyexceptionswereArea21andArea121whereonlyclippedindividualsweresampledandAreas20-1to20-4inAugustwhereonlyclippedindividualsweresampled Sampleswerepooled for analysis asoutlined inTable1withthelocationsofstatisticalareasoutlinedbyhttpswwwpacdfo-mpogccafm-gpmaps-cartesareas-secteursindex-enghtmlAdditionalcohosalmonsamples(bothclippedandunclipped)wereobtained froma troll test fisheryoperatingnearBrooksPeninsulaoff thenorthwest coastofVancouver Island a gillnet test fisherytargetingsockeyesalmon(Oncorhynchus nerka)nearRoundIslandinnorthernJohnstoneStraitandacommercialdemonstrationfreezertrollfisheryinthecentralcoastofBC(Figure1)
In2017forthenorthern(AreaF)freezertrollfisheryselectedfreezerboats(28offleet)wererequiredtokeepheadsofallcohosalmoncaughtwiththemarktype(adiposefinclippedornot)un-knownforanindividualheadUponlandingtheheadswerecountedand checked electronically for CWTs and randomly sampled to amaximumof50headsperdeliveryIfaCWTwasdetectedtheheadwassenttoacentralCWTheadrecoverylaboratoryinVancouverBCwheretheDNAsamplewassubsequentlytakenIfaCWTwassuccessfully recoveredanddecoded froman individualhead from
commercialrecreationalorFirstNationsfisheriesandagenotypewassuccessfullyobtainedforthe individualsample thenthegen-otypesofallof these individualswerepooled intoasinglemixed-stocksampleofknownoriginandknownage inorder toevaluateaccuracyof stock compositionsbyCUandpopulationFieldDNAsamplesweretakenonlyfromindividualswithnoCWTsdetectedInthenortherniceboattrollfisherytheclipstatusofthefishinthecatchwasknownSamplesof clippedcohosalmonwereobtainedfrom this fishery as an ancillary aspect of standard Fisheries andOceansCanadacontractcatchsamplingforCWTsOnlyclippedin-dividualswereexaminedthroughthisprogramwithsimilarsamplingprotocolsasoutlinedinthefreezerboatsamplingInparticularonlyclippedfishnotcontainingaCWTweresampledinthefieldthroughthisprogramwithheadscontainingaCWTsenttotheheadrecov-ery laboratory inVancouverClipped fish constituted16of thecatchinthefisherylargelyduetothewildoriginofthecatchsoitwasimportanttoobtainsamplesfromunclippedindividualsaswellSamplesofunclipped individualswereobtainedby sampling land-ingsforamaximumof25individualspervesselwithamaximumof100 individuals sampled per day Samples from theCentralCoastfreezertrolldemonstrationfisherywereobtainedviaasimilarpro-tocolasoutlinedforthenorthernfreezertrollsamplingTheoriginofthesamplesfromtherecreationalfisheryinBCincludedvolun-taryheadrecoveriesofadiposefin-clippedcohosalmonfromrecre-ationalfisheriesinBCaswellassomecreelsamplingofrecreationalcatches Samples from the recreational fisherywerederived fromclippedindividualsbuttheymaynothavebeenmarkedwithaCWTwhendeliveredtotheCWTheadrecoverylaboratoryThussamplesin2017wereobtainedfromall individualsthatwouldberoutinely
Fishery Reporting region Statistical area
Recreational JohnstoneStrait(JST) A11toA13
StraitofGeorgia-north(SOG-n) A14toA16
StraitofGeorgia-south(SOG-s) A17toA18A19-7toA19-12A28A29
StraitofGeorgiaJuandeFucaStrait(SOGJDF)
A19-1toA19-6A20-5
WestcoastVancouverIslandJuandeFucaStrait(WCVIJDF)
A20(exceptA20-5)
SouthwestVancouverIsland(SWVI21121)
A21A121
SouthwestVancouverIsland-inshore(SWVI-inshore)
A23A24
SouthwestVancouverIsland-offshore(SWVI-offshore)
A123A124
NorthwestVancouverIsland-inshore(NWVI-inshore)
A25toA27
NorthwestVancouverIsland-offshore(NWVI-offshore)
A125toA127
Test BrooksPeninsula A27A127A126
RoundIsland A12
Troll CentralCoast A6A7A8
TA B L E 1 emspReportingregionandDFOstatisticalareaswithinregionsforamalgamationofcatchsamplesLocationsofstatisticalareasareoutlinedonhttpswwwpacdfo-mpogccafm-gpmaps-cartesareas-secteursindex-enghtml
emspensp emsp | emsp5BEACHAM Et Al
processed under the CWT recovery program for coho salmon inBCInadditionsampleswerecollectedfromarecreationalfishery(clippedandunclipped individuals) fromHaidaGwaiinearLangaraIslandarecreational fisherynearRivers Inlet (Area8clippedandunclippedindividuals)inthecentralcoastthepreviouslynotediceboat northern troll fishery and direct creel sampling (clipped andunclippedindividuals)ofrecreationalfisheriesinsouthernBC
Monthly catch composition estimates were determined forclipped individuals from various fisheries in BC Seasonal catchcomposition estimateswere determined by recording the numberofindividualgenotypesavailablefromthefisherysamplingineachmonth andweighting individualmonthly samples by the recordedcatch in themonth Individualswere randomlychosen formonthsinwhich the number of available genotypeswas in excess of thenumberrequiredfortheseasonalsampleForexampleif60oftheannualcatchwasrecordedinonemonthand200genotypeswereavailablefromthefisherythenallindividualssampledwouldbein-cludedintheseasonalsamplebutcontributionsofindividualsfromothermonthswerescaledrelativetothemonthlycontributiontothe
seasonalcatchIfthecatchinoneothermonthwas20oftheannualcatchand150sampleswereavailablethen200(2060)=67genotypeswererandomlychosenfromtheavailable150genotypesforinclusionintotheseasonalsample
InthenorthernBCtrollfisherysamplerselectronicallycheckedclippedindividualsforthepresenceofaCWTandheadsfromthoseindividualscontainingaCWTweresubsequentlysenttothecentrallaboratory for CWT recovery and tissue sampling for subsequentgenotypingTissuesamplesfromclippedindividualswithnoCWTsweredirectlyprovidedforgenotypingInordertoestimatemonthlyandseasonalstockcompositionofclippedindividualsinthisfisherywepooledthegenotypesobtainedfromdirectfisherysamplingofclippedindividualswiththosesubsequentlysenttothecentrallabo-ratoryforCWTrecoveryinordertoprovideanappropriatepoolofclippedindividualstoestimatemonthlyandseasonalstockcompo-sitionsofclippedindividualsinthefisheryIneachsampleanalyzedover all BC fisheries thenumberof individuals identified viaPBTrelative to thenumberofgenotypes in the samplewas tabulatedandsummarizedoversamplefisheryandseason
F I G U R E 1 emspMapindicatinggeographiclocationsforfisherysamplingand20populationsforwhichparentage-basedtaggingwasappliedinestimationofstockcomposition
6emsp |emsp emspensp BEACHAM Et Al
22emsp|emspEscapement sample collection
In2016adiposefin-clippedjacks(age2years)weresampledfromeightpopulationswith573individualscollectedandgenotypedsuc-cessfullyJacksweredefinedonthebasisofbodysizeduringsam-plecollectionwith jacksvisiblysmallerthanage3-yearspawnersPopulations sampled were as follows Robertson Creek QuinsamRiver Puntledge River Big Qualicum River Goldstream RiverChilliwackRiverStaveRiverand InchCreekTheobjectiveofthesamplingwas to obtain a sample of known origin and known agebaseduponknowncollectionlocationinordertoevaluateaccuracyofassignmentsof the jacks tobothCUandpopulationunder theassumption of no straying among populations for the individualssampled
In 2017 escapements (non-broodstock hatchery and river re-turns clipped individualsonly)were sampledwhere samplingwasfeasibleEscapementsamplingfrom13populationswasconductedand1692 individualsweregenotypedEscapement sampleswerenot available from all 20 populations forwhich itwas possible toassignindividualsinthenon-broodstockescapementsviaPBTTheobjectiveofthesamplingwastoevaluatestrayingratesamongpop-ulationsandforthosehatcheries(ChilliwackRiver)whereparentalgenotypeswereassignedtospecificreleasegroupsprovide infor-mationon the relative rates of returnof different release groupsAs relative ratesof return for the releasegroupswereconsideredancillarytothemainfocusofthestudynoresultsfromthesereturnswereincludedinthecurrentstudy
23emsp|emspExploitation rate
WeestimatedexploitationrateofadultcohosalmoninBCfisheriesviabothCWTsandgeneticsExploitationrateforapopulationwasdefinedasadultcatch(adultcatch-escapement)ForCWTstheob-servednumberofCWTswascorrectedbyldquono-pinrdquotaglossratesandwasexpandedbythepopulationrsquostag-specificmarkingratesummedovertagcodesandexpandedagainbythesamplingrateforthefish-eryinordertoestimatecatchofhatchery-originindividualsTheob-servednumberofCWTsinescapementsamplingwasexpandedinasimilarmanner inordertoestimatethehatcherycontributiontothe escapement For genetics the seasonal kept catchof adiposefin-clippedindividualsinafisherywasmultipliedbyaseasonalstockcompositionestimateinordertoestimatepopulation-specificcatchTheabundanceofhatchery-originescapementwascalculatedastheestimatedescapementmultipliedbytheproportionofadiposefin-clippedadultsobserved in theescapement If partof the juvenileproductionwasnotadiposefinclippeduponreleasefromthehatch-erythenthehatcherycontributionstobothcatchandescapementwereunderestimatedbythismethod
24emsp|emspGenetic stock identification baseline
The initial baseline was outlined by Beacham et al (2017) andconsisted of 20242 individuals from 117 populations with the
distributionofpopulationsrangingfromsoutheastAlaskatoPugetSoundinWashingtonStateThebaselinehasbeensubsequentlyex-panded to include 40774 individuals from267populations rang-ingfromsoutheastAlaskatoOregonTheprimaryexpansionofthebaselineincludedasurveyofadditionalpopulationsinsouthernBCcoastalWashingtontheColumbiaRiverdrainageandOregonThefull baseline is outlined in Supporting Information Table S1 withthe populations fromBC arrangedbyCU andwithUnited StatesofAmerica(US)populationsarrangedbygeographic(reporting)re-gionBeachametal (2017)hadpreviouslydemonstratedastrongregionalpopulationstructureandreportedthatoverall117popula-tionsaverageself-assignmentaccuracytopopulation(weightedbypopulation sample size)was 855 for individualswith an assign-mentprobabilitylt050excludedand928forindividualswithanassignmentprobabilitylt085excludedHighlevelsofaccuracyforGSI assignment of individuals to specific populations when com-binedwithassignmentsviaPBTprovideapowerful technique forassignmentofindividualsofunknownorigintosomepopulations
25emsp|emspLibrary preparation and genotyping
ThedetailedprocedureforlibrarypreparationandgenotypingwasoutlinedbyBeachametal(2017)SummarizedbrieflyDNAwasex-tractedfromcohosalmontissuesamplesviaaChelexextractionandtheDNAconcentration normalized to40ngμlwith a Tecan LiHarobotApanelofprimersdesignedtoamplifyspecificsegmentsofDNAthatcontainedSNPsofinterestwasappliedtotheextractedDNAwithprimersequencesforeachampliconoutlinedbyBeachametal (2017SupportingInformationTableS2)TheinitialmultiplexPCR amplification of 304 target ampliconswas conductedwith acocktail of 2μl of normalizedDNA extract 5μl of 2X Ion Agriseqprimerpool2μlofIonAgriseqHiFimixand1μlofddH2O Thermal cyclingwasconductedin96-wellPCRplates(oneindividualperwell)withthefollowingconditionsforPCR99degCndash2min17cycles(99degCndash15s60degCndash4min)10degChold
FollowingtheinitialPCRasecondstepemployingathermalcy-clerwasconducted thatpartiallydigested theprimerson theam-pliconsandthereactionconductedwiththefollowingconditions50degCndash10min55degCndash10min60degCndash20min10degCholdA thirdandfinalstepemployingathermalcyclerwasinitiatedtoligatethebarcodes(768individualcodes)totheampliconsandwasconductedwiththefollowingconditions22degCndash30min70degCndash10min10degCholdLibrarieswerepurifiedbyadditionof225μlofAgencourtregAMPureregXPmagneticbeadstoeachlibrarytheplatewasplacedonamagnetic racksupernatantdiscardedandthebeadswashedtwice in70ethanolThepurified librarieswerethenelutedwith25 μloflowTEand20μlofthesupernatanttransferredtoafresh96-well trayNext eachof the768prepared librarieswaspooledintoasingletubeforprocessingontheIonChefreg(ThermoFisherScientific)Twotubesofpooled librarieswereprocessedconsecu-tivelyontheIonChefandthus1536individualswereprocessedon a single run of the Ion ChefOne tube of the pooled librarieswasloadedontoeachP1regchipv3andthusampliconsfrom768
emspensp emsp | emsp7BEACHAM Et Al
individualsweredistributedoneachP1chipwith1536individualsprocessedbetweentwochipsThechipswerethenloadedontotheIonTorrentProtonsequencerAfter thesequencingrunwascom-pletedcomparisonswiththereferencegenomeoftherainbowtrout(O mykiss)(Berthelotetal2014)supplementedwiththesequencescontaining theobservedcohosalmonSNPswereconductedwithProton softwareVariant Callerreg and SNP genotypes at the sitesspecified by the hotspot filewithin target regionswere called byVariantCallerThehotspotfilecontained304SNPsiteswithoneSNPscoredateachampliconGenotypesatallavailableSNPsforanindividualwereassembled toprovideamulti-locus individualgen-otype The species identification SNPOkiOts_120255‐113 (StarksClementoampGarza2016)andsex identificationSNPOts_SEXY3‐1 wereomittedfromsubsequentparentageandGSIanalysesleaving302SNPs
26emsp|emspIdentification of individuals
AsnotedpreviouslyPBTandGSIwereusedconcurrently toesti-mate fishery stock composition Initially PBT was used and theanalysiswas conductedwhere the genotypes of individuals to beidentifiedwerematched to the genotypes of prospective parents(COLONY JonesampWang 2010Wang 2016) If all individuals ina hatchery broodstock are sampled and subsequently genotypedthenalloffspringfromthebroodstockaregeneticallymarkedThegenotypes of individuals of unknown origin are statistically com-pared with the genotypes of potential parents and if a match ismadetheoffspringareassignedtotheparentsandthustheori-ginandageoftheindividualsaredeterminedParentageassignmentsoftwarewasutilizedtoassignoffspringtoparentsasCOLONYcanproduceassignmentswhen thegenotypeofoneof theparents ismissingeitherduetoamissingparentalsampleor failure topro-duceaparentalgenotypefromanexistingsampleGiventhatPBTassignments for20potential populationswereevaluated for eachfisherysampleCOLONYwasrunwithallbroodstocksampleddur-ing2014inputasasingleunitforanalysisoffisherysampleswithnodifferentiationamongpopulationsAlthoughtheCOLONYassump-tionofasinglepopulationintheparentpoolwasviolatedanalysisofknown-originsamplesindicatedthatveryhighlevelsofaccuracywereachievedinassignmentswhenpoolingofpotentialparentsincontributing populationswas conducted Two-parent assignmentswere accepted only when both assigned parents originated fromthesamepopulationotherwisetheindividualwaspassedtogeneticstockidentification(GSI)forpotentialassignmentTwo-parentandsingle-parentassignmentswereacceptedonlywhentheprobabilityofcorrectassignmentwasge085fortheparentpairotherwisetheindividualwaspassedtopotentialassignmentbyGSIAdditionallyforsingle-parentassignments tobeaccepted thePBTassignmenttopopulationhad tobepart of theCUassigned to the individualviaGSI Individuals forwhichnoprospectiveparentswere identi-fiedinthebroodstockwerepassedtoGSIforpotentialassignmentPolygamousmatingwasassumedfortheCOLONYanalysisSimplepairwisecomparisonsbetweenoffspringandpotentialparentswere
conductedThebaselineforindividualssampledinthe2016escape-ments (jacks)and2017fisheries includedallbroodstockssampledin2014as these individualsarepredominately threeyearsofage(Sandercock1991)Jacks inthe2017escapementwere identifiedvia body size and subsequent assignment to parents in the 2015hatcherybroodstocksIndividualswithmorethan120missinggeno-typeswereeliminatedfromfurtheranalysesAnestimatedgenotyp-ingerrorrateof1wasusedforCOLONYassignmentsPreviouslyBeachametal(2017)hadreportedthatanaveragegenotypingerrorrate of 107 (1220discrepancies in 114105 comparisons) or analleleerrorrateof053(1220discrepanciesin228210compari-sons)wasobservedoverthe302SNPsscoredTheparentpairout-putfilewasthebasicfileusedinsubsequentanalyses
Thesecondmethodof individual identification isGSI inwhichthegeneticprofilesofwholepopulationspotentiallycontributingtoamixed-stocksampleareusedtoestimatetheoriginofeachindivid-ualinthesample(RUBIASMoranampAnderson2018)ThisanalysiswasrestrictedtothoseindividualsunassignedviaCOLONYForeachsample individuals not assigned by COLONYwere then assignedwithRUBIASThepopulationposteriormeansfilewasthebasicfileusedforsubsequentanalysesandthisfilecontainedtheprobabilityofassignmentoftheindividualtoeachofthe267populationsinthebaselineStockcompositionwasestimatedthroughthecombinationoffilesgeneratedwithbothCOLONYandRUBIAS Individualsas-signedviaCOLONYwereassignedaprobabilityof100oforiginat-ingfromtheidentifiedpopulationwitha000probabilityassignedtoallotherpopulationsinthebaselineThislevelofassignmentac-curacyviaPBTwasobservedpreviouslyforcohosalmonwiththepanelofSNPsemployedinthecurrentstudy(Beachametal2017)ThesedatawerethencombinedwiththeprobabilityofassignmentforthoseindividualsunassignedviaCOLONYtoeachofthepopu-lationsinthebaselineviaRUBIASAfteraninitialburn-inof25000iterations the last1000 iterations from theMonteCarloMarkovchainfromRUBIASwereusedtoestimatetheoriginofindividualsandstockcompositionwiththemeanallocationtoeachpopulationinthebaselinedeterminedStandarddeviationsofestimatedstockcompositionswere alsodetermined from the last1000 iterationsfrom theMonteCarloMarkovchain Stock compositionbyCUorreportinggroupwasdeterminedbysummationofallocationstoallpopulations in the baseline that belonged to the CU or reportinggroupunderconsideration
3emsp |emspRESULTS
31emsp|emspEstimation of stock composition for known‐origin samples
Genotypeswere available from573 jacks sampled in2016acrossthreeCUs and eight populations in southernBC Estimated stockcompositionbyCUand reportinggroup for this combinedsamplewas accuratewith an error utilizingonlyGSI of le04byCU forthreeCUspresentinthesample(Table2)WithbothPBTandGSIutilized theaverageerrordeclinedtole02byCUfor threeCUs
8emsp |emsp emspensp BEACHAM Et Al
TAB
LE 2
emspAccuracyofregional(UnitedStates)andconservationunit(Canada)estimatedstockcomposition()foramixed-stocksampleof573known-origincohosalmonjackssampledin
2016and620coded-wiretaggedindividualssampledin2017fisheriesinBCStandarddeviationoftheestimateisinparentheses
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
SoutheastAlaska
00
00(00)
00(00)
90
68(11)
68(11)
AlsekRiver
00
00(00)
00(00)
00
00(00)
00(00)
LowerStikine
00
00(00)
00(00)
00
04(03)
04(03)
LowerNass
00
00(00)
00(00)
63
64(10)
62(10)
UpperNass
00
00(00)
00(00)
00
04(03)
04(04)
PortlandSound-Observatory
Inlet-PortlandCanal
00
00(00)
00(00)
00
00(00)
00(00)
SkeenaEstuary
00
00(00)
00(00)
00
00(00)
00(00)
LowerSkeena
00
00(00)
00(00)
16
12(05)
12(05)
MiddleSkeena
00
00(00)
00(00)
69
39(08)
39(09)
UpperSkeena
00
00(00)
00(00)
06
40(10)
40(09)
HaidaGwaiindashGrahamIslandLowlands
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-East
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-W
est
00
00(00)
00(00)
00
00(00)
00(00)
NorthernCoastalStreams
00
00(00)
00(00)
00
00(00)
00(01)
HecateStraitMainland
00
00(00)
00(00)
00
07(05)
06(04)
MusselndashKynoch
00
00(00)
00(00)
00
00(00)
00(00)
DouglasChannelndashKitimatArm
00
00(01)
00(01)
00
00(01)
00(01)
BellaCoolandashDeanRivers
00
00(00)
00(00)
00
00(00)
00(00)
RiversInlet
00
00(01)
00(00)
00
00(00)
00(00)
SmithInlet
00
00(00)
00(00)
00
00(00)
00(00)
SouthernCoastalStreamsndashQueen
CharlotteStraitndashJohnstoneStraitndash
SouthernFjords
00
00(01)
00(01)
00
00(02)
00(02)
HomathkondashKlinakliniRivers
00
00(00)
00(00)
00
00(00)
00(00)
GeorgiaStraitMainland
00
00(00)
00(00)
00
00(00)
00(00)
HoweSoundndashBurrardInlet
00
00(00)
00(00)
06
00(01)
00(00)
EastVancouverIslandndashGeorgiaStrait
351
345(20)
348(20)
131
134(14)
140(14)
EastVancouverIslandndashJohnstone
StraitndashSouthernFjords
00
00(00)
00(00)
00
00(01)
00(00)
NahwittiLowland
00
00(01)
00(00)
18
17(06)
17(06) (C
ontin
ues)
emspensp emsp | emsp9BEACHAM Et Al
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
WestVancouverIsland
115
115(14)
115(13)
135
134(14)
135(14)
Clayoquot
00
00(00)
00(00)
00
02(03)
02(02)
JuandeFucandashPachena
00
00(01)
00(01)
00
00(00)
00(00)
LowerFraser
534
539(21)
536(21)
185
187(15)
188(15)
Lillooet
00
00(00)
00(00)
00
03(02)
02(01)
FraserCanyon
00
00(00)
00(00)
00
00(00)
00(00)
InteriorFraser
00
00(00)
00(00)
00
00(00)
00(00)
LowerThompson
00
00(00)
00(00)
08
07(03)
08(04)
NorthThompson
00
00(00)
00(01)
00
00(00)
00(00)
SouthThompson
00
00(00)
00(00)
05
06(03)
05(03)
Boun
dary
Bay
00
00(00)
00(00)
00
00(00)
00(00)
NooksackRiver
00
01(02)
01(02)
26
09(09)
10(08)
SkagitRiver
00
00(01)
00(01)
05
23(08)
22(08)
NorthernPugetSound
00
00(00)
00(00)
39
58(12)
55(13)
Mid-PugetSound
00
00(00)
00(00)
52
62(11)
63(11)
SouthernPugetSound
00
00(00)
00(00)
10
09(10)
06(08)
JuandeFucaStrait
00
00(00)
00(00)
00
02(03)
01(03)
HoodCanal
00
00(00)
00(00)
44
16(05)
16(06)
CoastalWashington
00
00(00)
00(00)
60
59(10)
59(10)
ColumbiaRiver
00
00(00)
00(00)
32
32(07)
32(07)
Oregon
00
00(00)
00(00)
00
00(01)
00(00)
TAB
LE 2
(Continued)
10emsp |emsp emspensp BEACHAM Et Al
presentinthesample(Table2)Theaverageerrorfor45CUsorre-portinggroupsabsentinthesamplewas00bybothGSIandPBT-GSIBypopulation543(948)ofthejackswereassignedviaPBTwith 100 accuracywith respect to population of origin and age(Table3)Estimatedstockcompositionfor20populationswhereitwaspossibletousebothPBTandGSIinestimationofstockcompo-sitionwasaccurateFortheeightpopulationspresentinthesampleofjackstheaverageerrorutilizingGSIonlywas05perpopulationdecliningto02utilizingbothPBTandGSIundertheassumptionthatthecollectionlocationwasanaccuratereflectionofjackoriginTherewasanaverage00errorforthe12populationswithnorep-resentation inthesamplewithutilizationofeitherGSIorPBT-GSI(Table4)ThelargesterrorutilizingbothPBTandGSI(underestima-tionby06ofactualvalue)wasobservedintheGoldstreamRiverpopulationwhichalsodisplayedthe lowestpercentage (704)ofjacksidentifiedviaPBT(Table3)PBTwhencombinedwithGSIpro-ducedmoreaccurateestimatesofCU-specificandpopulation-spe-cificstockcompositionthanavailablewithonlyGSI
Of the 3054 adipose fin-clipped coho salmon samples re-ceivedforpotentialgenotypingfromacentrallaboratorywhere2017fisherysampleswereprocessedforpotentialCWTrecov-erygenotypeswereobtained for2533 (829)of the individ-uals processed There were 750 CWTs recovered from 3054headsor snouts examined and genotypeswereobtained from622 (829) of the individuals that carried a CWT GenotypesofcohosalmonwithoutCWTsbutsentfromtheCWTheadre-covery laboratorywereobtainedfrom1911of2304 individu-als (829) Release information associatedwith theCWTwasavailable for 620 of the 622CWTs associatedwith genotypedindividuals and these 620 individuals constituted a sample ofknown origin There were individuals from 20 regions or CUspresentinthissampleandforthese20regionsorCUstheav-erageerrorofestimationutilizingonlyGSIwas09perregionorCUand09forbothPBTandGSIcombined(Table2)CWTswererecoveredoriginatingfromsevenpopulationswhereitwaspossibletoincorporatebothGSIandPBTinestimationofstockcompositionFor these sevenpopulations theaverageerror inestimatedstockcompositionsperpopulationwas20withGSIand 03 with PBT-GSI (Table 4) There was an average 00errorforthe13populationswithnorepresentationinthesam-ple with utilization of either GSI or PBT-GSI (Table 4) Similartothesampleof jacksPBTwhencombinedwithGSIproducedmore accurate estimates of population-specific stock composi-tionthanavailablewithonlyGSI
32emsp|emspEvaluation of 2016 fishery sampling
Asbroodstocksamplingatselectedhatcheries insouthernBCdidnot commence until 2014 it was not possible to obtain PBT as-signments for any fishery samples collected in 2016 HoweverGSI analysis of individuals derived from recreational fishery sam-ples intheStraitofGeorgiaJuandeFucaStraitandoffthewestcoastofVancouverIslandinsouthernBCindicatedthattherewas
virtually no contribution from regions orCUsnorthofVancouverIsland (Supporting Information Table S2) The only exceptionwasin thenorthwestcoastofVancouver Islandboth inshoreandoff-shorewhereupto12ofthesamplewasidentifiedasoriginatingfromnorthernandcentralBCCUsUS-origincohosalmonwerees-timatedtohavecontributedapproximately70ofsummerrecrea-tionalsamplesderivedfromoffshorefisheriesalongthewestcoastofVancouverIslandaswellasintheentrancetoJuandeFucaStrait(A21A121)InmoreinshoreareasalongthewestcoastofVancouverIslandcohosalmonfromlocalCUswerethedominantcontributorstothefisheryForexamplethewestcoastofVancouverIslandCUcontributedapproximately80ofthe39-individualAugustsamplefromtheinshoresouthwesternVancouverIslandregionIntheStraitofGeorgia localCUswereagainthemaincontributorstotherec-reationalfisheryduringthesummerwiththeHoweSoundndashBurrardInletCUcontributingupto40insomesamplestheeastcoastofVancouverIslandCUuptoapproximately30andthelowerFraserRiverCUuptoapproximately40WhilethelowerFraserRiverCUwasanimportantcontributortotherecreationalfisheryintheStraitofGeorgiaandJuandeFucaStraititscontributiontofisheriesoffthewestcoastofVancouverIslandwaslimitedTheNooksackRiverregion innorthernWashingtonStatewasalso identifiedasacon-sistent contributor to recreational fishery samples in the Strait ofGeorgia insomecasescontributingupto20ofthesampleana-lyzed(SupportingInformationTableS2)
StockcompositionswereavailablefromtwotestfisheriesoneatrolltestfisherynearBrooksPeninsulabynorthwesternVancouverIslandandtheotheragillnettestfisheryinupperJohnstoneStraitIn the Brooks Peninsula fishery Vancouver IslandCUs comprised40ndash60ofthesampleswithaminorcontributionofFraserRiverCUs (lt10) The five interior Fraser River CUs of conservationconcern comprised00ndash33of the sampleswhereas themoreabundant lower Fraser River CUs comprised only 00ndash55 ofthesamples Individuals fromFraserRiverCUswerenotdetectedduringJulyinArea127theoffshoreportionofnorthernnorthwest-ernVancouverIslandbuttheyweredetectedinArea126themoresoutherlyoffshoreportionofnorthwesternVancouverIslandIntheJohnstoneStrait(RoundIsland)testfisheryindividualsfromFraserRiverCUswerescarcebutthefiveinteriorFraserRiverCUscom-prised30ndash35of the sampleswhereas the lowerFraserRiverCUscomprisedonly00ndash65ofthesamplesThemajorcontrib-utors to the samplesoriginated from theeast coastofVancouverIsland(24ndash45)andHoweSoundndashBurrardInletCUs(9ndash11)aswellascoastalWashington(8ndash15)(SupportingInformationTableS2)
ThefinalfisherysampledwasademonstrationcommercialtrollfisheryincentralcoastalBC(A6andA7)ContributionsfromlocalCUswerehigherinsamplesfrommoreinshoreareas(A7-7August212southernCUsandregions)withcontributionsfromsouth-ernCUshigherinthemoreseawardareas(A6-9September862southernCUsandregions)LikethewestcoastofVancouverIslandmigrating stocks were more likely to be found in more seawardlocations
emspensp emsp | emsp11BEACHAM Et Al
33emsp|emspEvaluation of 2017 fishery sampling
Onesourceoffisherysampleswasdirectfisherysamplingofindi-viduals(bothadiposefin-clippedandunclipped)wherethefocuswas not associated with potential CWT recovery In this casesamplingofunclippedindividualsfromthenorthern(AreaF)trollfishery indicated that initially (July 10ndash12) northern and centralBCCUscontributed23tothecatchsouthernBCCUs53andsouthern US regions 23 (Supporting Information Table S3)Stock composition changed markedly in samples collected July13ndash18 with Alaska contributing 12 northern and central BCCUs contributing 72 and southern BCCUs contributing 16withnosouthernUScontributionSamplescollectedduringJuly22ndash28revealedadominantcontributionfromnorthernandcen-tralBCCUs(96)andCUsfromthisregioncontributedbyfarthelargestportionof the catch in subsequent sampleswith south-ernBCCUsandsouthernUSregionslargelyabsent(SupportingInformation Table S3) The average contribution to the weeklysamples from July 22 onwardswas 17 southeast Alaska 16HaidaGwaiindashGrahamIslandLowlandsCU15DouglasChannelndashKitimatArmCU 12Hecate StraitMainlandCU 8NorthernCoastalStreamsCUand7LowerNassCUNotunexpectedlysampling of only clipped individuals from the fishery indicatedverysubstantialdifferencesinstockcompositionbetweenclippedandunclippedindividualsTheaveragecontributiontotheweeklysamples of clipped individuals from July 19 onwards was 32coastalWashington11mid-PugetSound10westVancouverIsland CU and 5 for each of northern Puget Sound and Juande Fuca Strait (Supporting Information Table S4) However theobservedcliprateinthefisherywas16sothecatchwasdomi-natedbyunclippedindividualsTheSeptember2ndash8samplefromthe Haida Gwaii recreational fishery near Langara displayed astockcompositionof42HaidaGwaiindashGrahamIslandLowlandsCU15HecateStraitMainlandCU10LowerSkeenaCUand8 Northern Coastal Streams CU similar to that observed inthe troll fisherywith the local CU (HaidaGwaiindashGraham IslandLowlands) inhigherproportionsaswouldbeexpectedinamoreinshorefishery(SupportingInformationTableS3)
Directsamplingof thecentralcoast troll fisheryrevealedthatonaverage theDouglasChannelndashKitimatArmCUwasthe largestcontributortothefishery(24)followedbytheNorthernCoastalStreamsCU(19)BellaCoolandashDeanRivers(18)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(10) (Supporting InformationTableS3)withallCUsproximaltothelocationofthefisherySamplesfromtherecreationalcentralcoastfisheryindicatedthattheBellaCoolandashDeanRiversCUwasthedominant contributor to the fishery (32) followedby theRiversInletCU(19)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(15)andDouglasChannelndashKitimatArmCU(7) (SupportingInformationTableS3)GiventhelocationofthefisherydominantcontributionsfromtheseCUsweretobeexpected
AsnotedpreviouslysampleswerealsoavailablefromvirtuallyallindividualssampledfromcommercialrecreationalandFirstNationsfisheriesthatwouldberoutinelyprocessedforCWTrecoveryinayearbyacentrallaboratoryIndividualheadsfromcommercialfish-eries sent to the central laboratoryhadbeenpreviously screenedelectronically for thepresenceofaCWTVoluntaryheadsamplesfromadiposefin-clippedindividualsfromrecreationalfisheriesweresenttothecentrallaboratorybutmanyoftheseindividualsdidnotcontainaCWTastherewasnopriorscreeningof these individu-als Stock composition of clipped individuals in the northern trollfishery has been previously outlined above Stock composition inthenorthern recreational fishery indicatedawidegeographicdis-tributionofcontributions rangingfromcoastalWashington (16)thewestcoastofVancouverIslandCU(10)theupperSkeenaCU(9)southeastAlaskaregion(8)mid-PugetSound(7)andeastcoast of Vancouver IslandndashStrait of Georgia CU (6) (SupportingInformation Table S4) Stock composition of clipped individuals inthecentralcoastrecreationalfisherywasdominatedbytheHoweSoundndashBurrardInletCU(27)andeastVancouverIslandndashStraitofGeorgiaCU(25)(SupportingInformationTableS4)butgiventhelowobservedclip rate in the fishery (014) therewasvery littlecatchthatcouldbeidentifiedashatcheryorigin
Directsamplingofrecreationalfisheries insouthernBCwhichincluded both clipped and unclipped individuals in the sample
Population Genotyped Assigned assignment
accuracy of population assignment
Robertson 66 66 1000 1000
Quinsam 69 68 986 1000
Puntledge 21 19 905 1000
BigQualicum 84 84 1000 1000
Goldstream 27 19 704 1000
Chilliwack 195 183 938 1000
Inch 91 85 934 1000
Stave 20 19 950 1000
Total 573 543 948 1000
TA B L E 3 emspPercentageofadiposefin-clippedjacks(age2years)sampledin2016andassignedviaparentageanalysistoparentsin2014broodstockpopulationsPopulationofsamplingwasassumedtobethepopulationoforiginofthejacksforevaluationofaccuracyofassignment
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
4emsp |emsp emspensp BEACHAM Et Al
2emsp |emspMETHODS
21emsp|emspFishery sample collection
Atotalof8006 individualsweregenotyped fromfisherysamplescollected in2016and2017 In2016sampleswerecollectedfromadiposefin-clippedandunclippedcohosalmonlandedintherecrea-tionalfisheryintheStraitofGeorgiaJuandeFucaStraitandalongthewestcoastofVancouverIslandTheonlyexceptionswereArea21andArea121whereonlyclippedindividualsweresampledandAreas20-1to20-4inAugustwhereonlyclippedindividualsweresampled Sampleswerepooled for analysis asoutlined inTable1withthelocationsofstatisticalareasoutlinedbyhttpswwwpacdfo-mpogccafm-gpmaps-cartesareas-secteursindex-enghtmlAdditionalcohosalmonsamples(bothclippedandunclipped)wereobtained froma troll test fisheryoperatingnearBrooksPeninsulaoff thenorthwest coastofVancouver Island a gillnet test fisherytargetingsockeyesalmon(Oncorhynchus nerka)nearRoundIslandinnorthernJohnstoneStraitandacommercialdemonstrationfreezertrollfisheryinthecentralcoastofBC(Figure1)
In2017forthenorthern(AreaF)freezertrollfisheryselectedfreezerboats(28offleet)wererequiredtokeepheadsofallcohosalmoncaughtwiththemarktype(adiposefinclippedornot)un-knownforanindividualheadUponlandingtheheadswerecountedand checked electronically for CWTs and randomly sampled to amaximumof50headsperdeliveryIfaCWTwasdetectedtheheadwassenttoacentralCWTheadrecoverylaboratoryinVancouverBCwheretheDNAsamplewassubsequentlytakenIfaCWTwassuccessfully recoveredanddecoded froman individualhead from
commercialrecreationalorFirstNationsfisheriesandagenotypewassuccessfullyobtainedforthe individualsample thenthegen-otypesofallof these individualswerepooled intoasinglemixed-stocksampleofknownoriginandknownage inorder toevaluateaccuracyof stock compositionsbyCUandpopulationFieldDNAsamplesweretakenonlyfromindividualswithnoCWTsdetectedInthenortherniceboattrollfisherytheclipstatusofthefishinthecatchwasknownSamplesof clippedcohosalmonwereobtainedfrom this fishery as an ancillary aspect of standard Fisheries andOceansCanadacontractcatchsamplingforCWTsOnlyclippedin-dividualswereexaminedthroughthisprogramwithsimilarsamplingprotocolsasoutlinedinthefreezerboatsamplingInparticularonlyclippedfishnotcontainingaCWTweresampledinthefieldthroughthisprogramwithheadscontainingaCWTsenttotheheadrecov-ery laboratory inVancouverClipped fish constituted16of thecatchinthefisherylargelyduetothewildoriginofthecatchsoitwasimportanttoobtainsamplesfromunclippedindividualsaswellSamplesofunclipped individualswereobtainedby sampling land-ingsforamaximumof25individualspervesselwithamaximumof100 individuals sampled per day Samples from theCentralCoastfreezertrolldemonstrationfisherywereobtainedviaasimilarpro-tocolasoutlinedforthenorthernfreezertrollsamplingTheoriginofthesamplesfromtherecreationalfisheryinBCincludedvolun-taryheadrecoveriesofadiposefin-clippedcohosalmonfromrecre-ationalfisheriesinBCaswellassomecreelsamplingofrecreationalcatches Samples from the recreational fisherywerederived fromclippedindividualsbuttheymaynothavebeenmarkedwithaCWTwhendeliveredtotheCWTheadrecoverylaboratoryThussamplesin2017wereobtainedfromall individualsthatwouldberoutinely
Fishery Reporting region Statistical area
Recreational JohnstoneStrait(JST) A11toA13
StraitofGeorgia-north(SOG-n) A14toA16
StraitofGeorgia-south(SOG-s) A17toA18A19-7toA19-12A28A29
StraitofGeorgiaJuandeFucaStrait(SOGJDF)
A19-1toA19-6A20-5
WestcoastVancouverIslandJuandeFucaStrait(WCVIJDF)
A20(exceptA20-5)
SouthwestVancouverIsland(SWVI21121)
A21A121
SouthwestVancouverIsland-inshore(SWVI-inshore)
A23A24
SouthwestVancouverIsland-offshore(SWVI-offshore)
A123A124
NorthwestVancouverIsland-inshore(NWVI-inshore)
A25toA27
NorthwestVancouverIsland-offshore(NWVI-offshore)
A125toA127
Test BrooksPeninsula A27A127A126
RoundIsland A12
Troll CentralCoast A6A7A8
TA B L E 1 emspReportingregionandDFOstatisticalareaswithinregionsforamalgamationofcatchsamplesLocationsofstatisticalareasareoutlinedonhttpswwwpacdfo-mpogccafm-gpmaps-cartesareas-secteursindex-enghtml
emspensp emsp | emsp5BEACHAM Et Al
processed under the CWT recovery program for coho salmon inBCInadditionsampleswerecollectedfromarecreationalfishery(clippedandunclipped individuals) fromHaidaGwaiinearLangaraIslandarecreational fisherynearRivers Inlet (Area8clippedandunclippedindividuals)inthecentralcoastthepreviouslynotediceboat northern troll fishery and direct creel sampling (clipped andunclippedindividuals)ofrecreationalfisheriesinsouthernBC
Monthly catch composition estimates were determined forclipped individuals from various fisheries in BC Seasonal catchcomposition estimateswere determined by recording the numberofindividualgenotypesavailablefromthefisherysamplingineachmonth andweighting individualmonthly samples by the recordedcatch in themonth Individualswere randomlychosen formonthsinwhich the number of available genotypeswas in excess of thenumberrequiredfortheseasonalsampleForexampleif60oftheannualcatchwasrecordedinonemonthand200genotypeswereavailablefromthefisherythenallindividualssampledwouldbein-cludedintheseasonalsamplebutcontributionsofindividualsfromothermonthswerescaledrelativetothemonthlycontributiontothe
seasonalcatchIfthecatchinoneothermonthwas20oftheannualcatchand150sampleswereavailablethen200(2060)=67genotypeswererandomlychosenfromtheavailable150genotypesforinclusionintotheseasonalsample
InthenorthernBCtrollfisherysamplerselectronicallycheckedclippedindividualsforthepresenceofaCWTandheadsfromthoseindividualscontainingaCWTweresubsequentlysenttothecentrallaboratory for CWT recovery and tissue sampling for subsequentgenotypingTissuesamplesfromclippedindividualswithnoCWTsweredirectlyprovidedforgenotypingInordertoestimatemonthlyandseasonalstockcompositionofclippedindividualsinthisfisherywepooledthegenotypesobtainedfromdirectfisherysamplingofclippedindividualswiththosesubsequentlysenttothecentrallabo-ratoryforCWTrecoveryinordertoprovideanappropriatepoolofclippedindividualstoestimatemonthlyandseasonalstockcompo-sitionsofclippedindividualsinthefisheryIneachsampleanalyzedover all BC fisheries thenumberof individuals identified viaPBTrelative to thenumberofgenotypes in the samplewas tabulatedandsummarizedoversamplefisheryandseason
F I G U R E 1 emspMapindicatinggeographiclocationsforfisherysamplingand20populationsforwhichparentage-basedtaggingwasappliedinestimationofstockcomposition
6emsp |emsp emspensp BEACHAM Et Al
22emsp|emspEscapement sample collection
In2016adiposefin-clippedjacks(age2years)weresampledfromeightpopulationswith573individualscollectedandgenotypedsuc-cessfullyJacksweredefinedonthebasisofbodysizeduringsam-plecollectionwith jacksvisiblysmallerthanage3-yearspawnersPopulations sampled were as follows Robertson Creek QuinsamRiver Puntledge River Big Qualicum River Goldstream RiverChilliwackRiverStaveRiverand InchCreekTheobjectiveofthesamplingwas to obtain a sample of known origin and known agebaseduponknowncollectionlocationinordertoevaluateaccuracyofassignmentsof the jacks tobothCUandpopulationunder theassumption of no straying among populations for the individualssampled
In 2017 escapements (non-broodstock hatchery and river re-turns clipped individualsonly)were sampledwhere samplingwasfeasibleEscapementsamplingfrom13populationswasconductedand1692 individualsweregenotypedEscapement sampleswerenot available from all 20 populations forwhich itwas possible toassignindividualsinthenon-broodstockescapementsviaPBTTheobjectiveofthesamplingwastoevaluatestrayingratesamongpop-ulationsandforthosehatcheries(ChilliwackRiver)whereparentalgenotypeswereassignedtospecificreleasegroupsprovide infor-mationon the relative rates of returnof different release groupsAs relative ratesof return for the releasegroupswereconsideredancillarytothemainfocusofthestudynoresultsfromthesereturnswereincludedinthecurrentstudy
23emsp|emspExploitation rate
WeestimatedexploitationrateofadultcohosalmoninBCfisheriesviabothCWTsandgeneticsExploitationrateforapopulationwasdefinedasadultcatch(adultcatch-escapement)ForCWTstheob-servednumberofCWTswascorrectedbyldquono-pinrdquotaglossratesandwasexpandedbythepopulationrsquostag-specificmarkingratesummedovertagcodesandexpandedagainbythesamplingrateforthefish-eryinordertoestimatecatchofhatchery-originindividualsTheob-servednumberofCWTsinescapementsamplingwasexpandedinasimilarmanner inordertoestimatethehatcherycontributiontothe escapement For genetics the seasonal kept catchof adiposefin-clippedindividualsinafisherywasmultipliedbyaseasonalstockcompositionestimateinordertoestimatepopulation-specificcatchTheabundanceofhatchery-originescapementwascalculatedastheestimatedescapementmultipliedbytheproportionofadiposefin-clippedadultsobserved in theescapement If partof the juvenileproductionwasnotadiposefinclippeduponreleasefromthehatch-erythenthehatcherycontributionstobothcatchandescapementwereunderestimatedbythismethod
24emsp|emspGenetic stock identification baseline
The initial baseline was outlined by Beacham et al (2017) andconsisted of 20242 individuals from 117 populations with the
distributionofpopulationsrangingfromsoutheastAlaskatoPugetSoundinWashingtonStateThebaselinehasbeensubsequentlyex-panded to include 40774 individuals from267populations rang-ingfromsoutheastAlaskatoOregonTheprimaryexpansionofthebaselineincludedasurveyofadditionalpopulationsinsouthernBCcoastalWashingtontheColumbiaRiverdrainageandOregonThefull baseline is outlined in Supporting Information Table S1 withthe populations fromBC arrangedbyCU andwithUnited StatesofAmerica(US)populationsarrangedbygeographic(reporting)re-gionBeachametal (2017)hadpreviouslydemonstratedastrongregionalpopulationstructureandreportedthatoverall117popula-tionsaverageself-assignmentaccuracytopopulation(weightedbypopulation sample size)was 855 for individualswith an assign-mentprobabilitylt050excludedand928forindividualswithanassignmentprobabilitylt085excludedHighlevelsofaccuracyforGSI assignment of individuals to specific populations when com-binedwithassignmentsviaPBTprovideapowerful technique forassignmentofindividualsofunknownorigintosomepopulations
25emsp|emspLibrary preparation and genotyping
ThedetailedprocedureforlibrarypreparationandgenotypingwasoutlinedbyBeachametal(2017)SummarizedbrieflyDNAwasex-tractedfromcohosalmontissuesamplesviaaChelexextractionandtheDNAconcentration normalized to40ngμlwith a Tecan LiHarobotApanelofprimersdesignedtoamplifyspecificsegmentsofDNAthatcontainedSNPsofinterestwasappliedtotheextractedDNAwithprimersequencesforeachampliconoutlinedbyBeachametal (2017SupportingInformationTableS2)TheinitialmultiplexPCR amplification of 304 target ampliconswas conductedwith acocktail of 2μl of normalizedDNA extract 5μl of 2X Ion Agriseqprimerpool2μlofIonAgriseqHiFimixand1μlofddH2O Thermal cyclingwasconductedin96-wellPCRplates(oneindividualperwell)withthefollowingconditionsforPCR99degCndash2min17cycles(99degCndash15s60degCndash4min)10degChold
FollowingtheinitialPCRasecondstepemployingathermalcy-clerwasconducted thatpartiallydigested theprimerson theam-pliconsandthereactionconductedwiththefollowingconditions50degCndash10min55degCndash10min60degCndash20min10degCholdA thirdandfinalstepemployingathermalcyclerwasinitiatedtoligatethebarcodes(768individualcodes)totheampliconsandwasconductedwiththefollowingconditions22degCndash30min70degCndash10min10degCholdLibrarieswerepurifiedbyadditionof225μlofAgencourtregAMPureregXPmagneticbeadstoeachlibrarytheplatewasplacedonamagnetic racksupernatantdiscardedandthebeadswashedtwice in70ethanolThepurified librarieswerethenelutedwith25 μloflowTEand20μlofthesupernatanttransferredtoafresh96-well trayNext eachof the768prepared librarieswaspooledintoasingletubeforprocessingontheIonChefreg(ThermoFisherScientific)Twotubesofpooled librarieswereprocessedconsecu-tivelyontheIonChefandthus1536individualswereprocessedon a single run of the Ion ChefOne tube of the pooled librarieswasloadedontoeachP1regchipv3andthusampliconsfrom768
emspensp emsp | emsp7BEACHAM Et Al
individualsweredistributedoneachP1chipwith1536individualsprocessedbetweentwochipsThechipswerethenloadedontotheIonTorrentProtonsequencerAfter thesequencingrunwascom-pletedcomparisonswiththereferencegenomeoftherainbowtrout(O mykiss)(Berthelotetal2014)supplementedwiththesequencescontaining theobservedcohosalmonSNPswereconductedwithProton softwareVariant Callerreg and SNP genotypes at the sitesspecified by the hotspot filewithin target regionswere called byVariantCallerThehotspotfilecontained304SNPsiteswithoneSNPscoredateachampliconGenotypesatallavailableSNPsforanindividualwereassembled toprovideamulti-locus individualgen-otype The species identification SNPOkiOts_120255‐113 (StarksClementoampGarza2016)andsex identificationSNPOts_SEXY3‐1 wereomittedfromsubsequentparentageandGSIanalysesleaving302SNPs
26emsp|emspIdentification of individuals
AsnotedpreviouslyPBTandGSIwereusedconcurrently toesti-mate fishery stock composition Initially PBT was used and theanalysiswas conductedwhere the genotypes of individuals to beidentifiedwerematched to the genotypes of prospective parents(COLONY JonesampWang 2010Wang 2016) If all individuals ina hatchery broodstock are sampled and subsequently genotypedthenalloffspringfromthebroodstockaregeneticallymarkedThegenotypes of individuals of unknown origin are statistically com-pared with the genotypes of potential parents and if a match ismadetheoffspringareassignedtotheparentsandthustheori-ginandageoftheindividualsaredeterminedParentageassignmentsoftwarewasutilizedtoassignoffspringtoparentsasCOLONYcanproduceassignmentswhen thegenotypeofoneof theparents ismissingeitherduetoamissingparentalsampleor failure topro-duceaparentalgenotypefromanexistingsampleGiventhatPBTassignments for20potential populationswereevaluated for eachfisherysampleCOLONYwasrunwithallbroodstocksampleddur-ing2014inputasasingleunitforanalysisoffisherysampleswithnodifferentiationamongpopulationsAlthoughtheCOLONYassump-tionofasinglepopulationintheparentpoolwasviolatedanalysisofknown-originsamplesindicatedthatveryhighlevelsofaccuracywereachievedinassignmentswhenpoolingofpotentialparentsincontributing populationswas conducted Two-parent assignmentswere accepted only when both assigned parents originated fromthesamepopulationotherwisetheindividualwaspassedtogeneticstockidentification(GSI)forpotentialassignmentTwo-parentandsingle-parentassignmentswereacceptedonlywhentheprobabilityofcorrectassignmentwasge085fortheparentpairotherwisetheindividualwaspassedtopotentialassignmentbyGSIAdditionallyforsingle-parentassignments tobeaccepted thePBTassignmenttopopulationhad tobepart of theCUassigned to the individualviaGSI Individuals forwhichnoprospectiveparentswere identi-fiedinthebroodstockwerepassedtoGSIforpotentialassignmentPolygamousmatingwasassumedfortheCOLONYanalysisSimplepairwisecomparisonsbetweenoffspringandpotentialparentswere
conductedThebaselineforindividualssampledinthe2016escape-ments (jacks)and2017fisheries includedallbroodstockssampledin2014as these individualsarepredominately threeyearsofage(Sandercock1991)Jacks inthe2017escapementwere identifiedvia body size and subsequent assignment to parents in the 2015hatcherybroodstocksIndividualswithmorethan120missinggeno-typeswereeliminatedfromfurtheranalysesAnestimatedgenotyp-ingerrorrateof1wasusedforCOLONYassignmentsPreviouslyBeachametal(2017)hadreportedthatanaveragegenotypingerrorrate of 107 (1220discrepancies in 114105 comparisons) or analleleerrorrateof053(1220discrepanciesin228210compari-sons)wasobservedoverthe302SNPsscoredTheparentpairout-putfilewasthebasicfileusedinsubsequentanalyses
Thesecondmethodof individual identification isGSI inwhichthegeneticprofilesofwholepopulationspotentiallycontributingtoamixed-stocksampleareusedtoestimatetheoriginofeachindivid-ualinthesample(RUBIASMoranampAnderson2018)ThisanalysiswasrestrictedtothoseindividualsunassignedviaCOLONYForeachsample individuals not assigned by COLONYwere then assignedwithRUBIASThepopulationposteriormeansfilewasthebasicfileusedforsubsequentanalysesandthisfilecontainedtheprobabilityofassignmentoftheindividualtoeachofthe267populationsinthebaselineStockcompositionwasestimatedthroughthecombinationoffilesgeneratedwithbothCOLONYandRUBIAS Individualsas-signedviaCOLONYwereassignedaprobabilityof100oforiginat-ingfromtheidentifiedpopulationwitha000probabilityassignedtoallotherpopulationsinthebaselineThislevelofassignmentac-curacyviaPBTwasobservedpreviouslyforcohosalmonwiththepanelofSNPsemployedinthecurrentstudy(Beachametal2017)ThesedatawerethencombinedwiththeprobabilityofassignmentforthoseindividualsunassignedviaCOLONYtoeachofthepopu-lationsinthebaselineviaRUBIASAfteraninitialburn-inof25000iterations the last1000 iterations from theMonteCarloMarkovchainfromRUBIASwereusedtoestimatetheoriginofindividualsandstockcompositionwiththemeanallocationtoeachpopulationinthebaselinedeterminedStandarddeviationsofestimatedstockcompositionswere alsodetermined from the last1000 iterationsfrom theMonteCarloMarkovchain Stock compositionbyCUorreportinggroupwasdeterminedbysummationofallocationstoallpopulations in the baseline that belonged to the CU or reportinggroupunderconsideration
3emsp |emspRESULTS
31emsp|emspEstimation of stock composition for known‐origin samples
Genotypeswere available from573 jacks sampled in2016acrossthreeCUs and eight populations in southernBC Estimated stockcompositionbyCUand reportinggroup for this combinedsamplewas accuratewith an error utilizingonlyGSI of le04byCU forthreeCUspresentinthesample(Table2)WithbothPBTandGSIutilized theaverageerrordeclinedtole02byCUfor threeCUs
8emsp |emsp emspensp BEACHAM Et Al
TAB
LE 2
emspAccuracyofregional(UnitedStates)andconservationunit(Canada)estimatedstockcomposition()foramixed-stocksampleof573known-origincohosalmonjackssampledin
2016and620coded-wiretaggedindividualssampledin2017fisheriesinBCStandarddeviationoftheestimateisinparentheses
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
SoutheastAlaska
00
00(00)
00(00)
90
68(11)
68(11)
AlsekRiver
00
00(00)
00(00)
00
00(00)
00(00)
LowerStikine
00
00(00)
00(00)
00
04(03)
04(03)
LowerNass
00
00(00)
00(00)
63
64(10)
62(10)
UpperNass
00
00(00)
00(00)
00
04(03)
04(04)
PortlandSound-Observatory
Inlet-PortlandCanal
00
00(00)
00(00)
00
00(00)
00(00)
SkeenaEstuary
00
00(00)
00(00)
00
00(00)
00(00)
LowerSkeena
00
00(00)
00(00)
16
12(05)
12(05)
MiddleSkeena
00
00(00)
00(00)
69
39(08)
39(09)
UpperSkeena
00
00(00)
00(00)
06
40(10)
40(09)
HaidaGwaiindashGrahamIslandLowlands
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-East
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-W
est
00
00(00)
00(00)
00
00(00)
00(00)
NorthernCoastalStreams
00
00(00)
00(00)
00
00(00)
00(01)
HecateStraitMainland
00
00(00)
00(00)
00
07(05)
06(04)
MusselndashKynoch
00
00(00)
00(00)
00
00(00)
00(00)
DouglasChannelndashKitimatArm
00
00(01)
00(01)
00
00(01)
00(01)
BellaCoolandashDeanRivers
00
00(00)
00(00)
00
00(00)
00(00)
RiversInlet
00
00(01)
00(00)
00
00(00)
00(00)
SmithInlet
00
00(00)
00(00)
00
00(00)
00(00)
SouthernCoastalStreamsndashQueen
CharlotteStraitndashJohnstoneStraitndash
SouthernFjords
00
00(01)
00(01)
00
00(02)
00(02)
HomathkondashKlinakliniRivers
00
00(00)
00(00)
00
00(00)
00(00)
GeorgiaStraitMainland
00
00(00)
00(00)
00
00(00)
00(00)
HoweSoundndashBurrardInlet
00
00(00)
00(00)
06
00(01)
00(00)
EastVancouverIslandndashGeorgiaStrait
351
345(20)
348(20)
131
134(14)
140(14)
EastVancouverIslandndashJohnstone
StraitndashSouthernFjords
00
00(00)
00(00)
00
00(01)
00(00)
NahwittiLowland
00
00(01)
00(00)
18
17(06)
17(06) (C
ontin
ues)
emspensp emsp | emsp9BEACHAM Et Al
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
WestVancouverIsland
115
115(14)
115(13)
135
134(14)
135(14)
Clayoquot
00
00(00)
00(00)
00
02(03)
02(02)
JuandeFucandashPachena
00
00(01)
00(01)
00
00(00)
00(00)
LowerFraser
534
539(21)
536(21)
185
187(15)
188(15)
Lillooet
00
00(00)
00(00)
00
03(02)
02(01)
FraserCanyon
00
00(00)
00(00)
00
00(00)
00(00)
InteriorFraser
00
00(00)
00(00)
00
00(00)
00(00)
LowerThompson
00
00(00)
00(00)
08
07(03)
08(04)
NorthThompson
00
00(00)
00(01)
00
00(00)
00(00)
SouthThompson
00
00(00)
00(00)
05
06(03)
05(03)
Boun
dary
Bay
00
00(00)
00(00)
00
00(00)
00(00)
NooksackRiver
00
01(02)
01(02)
26
09(09)
10(08)
SkagitRiver
00
00(01)
00(01)
05
23(08)
22(08)
NorthernPugetSound
00
00(00)
00(00)
39
58(12)
55(13)
Mid-PugetSound
00
00(00)
00(00)
52
62(11)
63(11)
SouthernPugetSound
00
00(00)
00(00)
10
09(10)
06(08)
JuandeFucaStrait
00
00(00)
00(00)
00
02(03)
01(03)
HoodCanal
00
00(00)
00(00)
44
16(05)
16(06)
CoastalWashington
00
00(00)
00(00)
60
59(10)
59(10)
ColumbiaRiver
00
00(00)
00(00)
32
32(07)
32(07)
Oregon
00
00(00)
00(00)
00
00(01)
00(00)
TAB
LE 2
(Continued)
10emsp |emsp emspensp BEACHAM Et Al
presentinthesample(Table2)Theaverageerrorfor45CUsorre-portinggroupsabsentinthesamplewas00bybothGSIandPBT-GSIBypopulation543(948)ofthejackswereassignedviaPBTwith 100 accuracywith respect to population of origin and age(Table3)Estimatedstockcompositionfor20populationswhereitwaspossibletousebothPBTandGSIinestimationofstockcompo-sitionwasaccurateFortheeightpopulationspresentinthesampleofjackstheaverageerrorutilizingGSIonlywas05perpopulationdecliningto02utilizingbothPBTandGSIundertheassumptionthatthecollectionlocationwasanaccuratereflectionofjackoriginTherewasanaverage00errorforthe12populationswithnorep-resentation inthesamplewithutilizationofeitherGSIorPBT-GSI(Table4)ThelargesterrorutilizingbothPBTandGSI(underestima-tionby06ofactualvalue)wasobservedintheGoldstreamRiverpopulationwhichalsodisplayedthe lowestpercentage (704)ofjacksidentifiedviaPBT(Table3)PBTwhencombinedwithGSIpro-ducedmoreaccurateestimatesofCU-specificandpopulation-spe-cificstockcompositionthanavailablewithonlyGSI
Of the 3054 adipose fin-clipped coho salmon samples re-ceivedforpotentialgenotypingfromacentrallaboratorywhere2017fisherysampleswereprocessedforpotentialCWTrecov-erygenotypeswereobtained for2533 (829)of the individ-uals processed There were 750 CWTs recovered from 3054headsor snouts examined and genotypeswereobtained from622 (829) of the individuals that carried a CWT GenotypesofcohosalmonwithoutCWTsbutsentfromtheCWTheadre-covery laboratorywereobtainedfrom1911of2304 individu-als (829) Release information associatedwith theCWTwasavailable for 620 of the 622CWTs associatedwith genotypedindividuals and these 620 individuals constituted a sample ofknown origin There were individuals from 20 regions or CUspresentinthissampleandforthese20regionsorCUstheav-erageerrorofestimationutilizingonlyGSIwas09perregionorCUand09forbothPBTandGSIcombined(Table2)CWTswererecoveredoriginatingfromsevenpopulationswhereitwaspossibletoincorporatebothGSIandPBTinestimationofstockcompositionFor these sevenpopulations theaverageerror inestimatedstockcompositionsperpopulationwas20withGSIand 03 with PBT-GSI (Table 4) There was an average 00errorforthe13populationswithnorepresentationinthesam-ple with utilization of either GSI or PBT-GSI (Table 4) Similartothesampleof jacksPBTwhencombinedwithGSIproducedmore accurate estimates of population-specific stock composi-tionthanavailablewithonlyGSI
32emsp|emspEvaluation of 2016 fishery sampling
Asbroodstocksamplingatselectedhatcheries insouthernBCdidnot commence until 2014 it was not possible to obtain PBT as-signments for any fishery samples collected in 2016 HoweverGSI analysis of individuals derived from recreational fishery sam-ples intheStraitofGeorgiaJuandeFucaStraitandoffthewestcoastofVancouverIslandinsouthernBCindicatedthattherewas
virtually no contribution from regions orCUsnorthofVancouverIsland (Supporting Information Table S2) The only exceptionwasin thenorthwestcoastofVancouver Islandboth inshoreandoff-shorewhereupto12ofthesamplewasidentifiedasoriginatingfromnorthernandcentralBCCUsUS-origincohosalmonwerees-timatedtohavecontributedapproximately70ofsummerrecrea-tionalsamplesderivedfromoffshorefisheriesalongthewestcoastofVancouverIslandaswellasintheentrancetoJuandeFucaStrait(A21A121)InmoreinshoreareasalongthewestcoastofVancouverIslandcohosalmonfromlocalCUswerethedominantcontributorstothefisheryForexamplethewestcoastofVancouverIslandCUcontributedapproximately80ofthe39-individualAugustsamplefromtheinshoresouthwesternVancouverIslandregionIntheStraitofGeorgia localCUswereagainthemaincontributorstotherec-reationalfisheryduringthesummerwiththeHoweSoundndashBurrardInletCUcontributingupto40insomesamplestheeastcoastofVancouverIslandCUuptoapproximately30andthelowerFraserRiverCUuptoapproximately40WhilethelowerFraserRiverCUwasanimportantcontributortotherecreationalfisheryintheStraitofGeorgiaandJuandeFucaStraititscontributiontofisheriesoffthewestcoastofVancouverIslandwaslimitedTheNooksackRiverregion innorthernWashingtonStatewasalso identifiedasacon-sistent contributor to recreational fishery samples in the Strait ofGeorgia insomecasescontributingupto20ofthesampleana-lyzed(SupportingInformationTableS2)
StockcompositionswereavailablefromtwotestfisheriesoneatrolltestfisherynearBrooksPeninsulabynorthwesternVancouverIslandandtheotheragillnettestfisheryinupperJohnstoneStraitIn the Brooks Peninsula fishery Vancouver IslandCUs comprised40ndash60ofthesampleswithaminorcontributionofFraserRiverCUs (lt10) The five interior Fraser River CUs of conservationconcern comprised00ndash33of the sampleswhereas themoreabundant lower Fraser River CUs comprised only 00ndash55 ofthesamples Individuals fromFraserRiverCUswerenotdetectedduringJulyinArea127theoffshoreportionofnorthernnorthwest-ernVancouverIslandbuttheyweredetectedinArea126themoresoutherlyoffshoreportionofnorthwesternVancouverIslandIntheJohnstoneStrait(RoundIsland)testfisheryindividualsfromFraserRiverCUswerescarcebutthefiveinteriorFraserRiverCUscom-prised30ndash35of the sampleswhereas the lowerFraserRiverCUscomprisedonly00ndash65ofthesamplesThemajorcontrib-utors to the samplesoriginated from theeast coastofVancouverIsland(24ndash45)andHoweSoundndashBurrardInletCUs(9ndash11)aswellascoastalWashington(8ndash15)(SupportingInformationTableS2)
ThefinalfisherysampledwasademonstrationcommercialtrollfisheryincentralcoastalBC(A6andA7)ContributionsfromlocalCUswerehigherinsamplesfrommoreinshoreareas(A7-7August212southernCUsandregions)withcontributionsfromsouth-ernCUshigherinthemoreseawardareas(A6-9September862southernCUsandregions)LikethewestcoastofVancouverIslandmigrating stocks were more likely to be found in more seawardlocations
emspensp emsp | emsp11BEACHAM Et Al
33emsp|emspEvaluation of 2017 fishery sampling
Onesourceoffisherysampleswasdirectfisherysamplingofindi-viduals(bothadiposefin-clippedandunclipped)wherethefocuswas not associated with potential CWT recovery In this casesamplingofunclippedindividualsfromthenorthern(AreaF)trollfishery indicated that initially (July 10ndash12) northern and centralBCCUscontributed23tothecatchsouthernBCCUs53andsouthern US regions 23 (Supporting Information Table S3)Stock composition changed markedly in samples collected July13ndash18 with Alaska contributing 12 northern and central BCCUs contributing 72 and southern BCCUs contributing 16withnosouthernUScontributionSamplescollectedduringJuly22ndash28revealedadominantcontributionfromnorthernandcen-tralBCCUs(96)andCUsfromthisregioncontributedbyfarthelargestportionof the catch in subsequent sampleswith south-ernBCCUsandsouthernUSregionslargelyabsent(SupportingInformation Table S3) The average contribution to the weeklysamples from July 22 onwardswas 17 southeast Alaska 16HaidaGwaiindashGrahamIslandLowlandsCU15DouglasChannelndashKitimatArmCU 12Hecate StraitMainlandCU 8NorthernCoastalStreamsCUand7LowerNassCUNotunexpectedlysampling of only clipped individuals from the fishery indicatedverysubstantialdifferencesinstockcompositionbetweenclippedandunclippedindividualsTheaveragecontributiontotheweeklysamples of clipped individuals from July 19 onwards was 32coastalWashington11mid-PugetSound10westVancouverIsland CU and 5 for each of northern Puget Sound and Juande Fuca Strait (Supporting Information Table S4) However theobservedcliprateinthefisherywas16sothecatchwasdomi-natedbyunclippedindividualsTheSeptember2ndash8samplefromthe Haida Gwaii recreational fishery near Langara displayed astockcompositionof42HaidaGwaiindashGrahamIslandLowlandsCU15HecateStraitMainlandCU10LowerSkeenaCUand8 Northern Coastal Streams CU similar to that observed inthe troll fisherywith the local CU (HaidaGwaiindashGraham IslandLowlands) inhigherproportionsaswouldbeexpectedinamoreinshorefishery(SupportingInformationTableS3)
Directsamplingof thecentralcoast troll fisheryrevealedthatonaverage theDouglasChannelndashKitimatArmCUwasthe largestcontributortothefishery(24)followedbytheNorthernCoastalStreamsCU(19)BellaCoolandashDeanRivers(18)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(10) (Supporting InformationTableS3)withallCUsproximaltothelocationofthefisherySamplesfromtherecreationalcentralcoastfisheryindicatedthattheBellaCoolandashDeanRiversCUwasthedominant contributor to the fishery (32) followedby theRiversInletCU(19)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(15)andDouglasChannelndashKitimatArmCU(7) (SupportingInformationTableS3)GiventhelocationofthefisherydominantcontributionsfromtheseCUsweretobeexpected
AsnotedpreviouslysampleswerealsoavailablefromvirtuallyallindividualssampledfromcommercialrecreationalandFirstNationsfisheriesthatwouldberoutinelyprocessedforCWTrecoveryinayearbyacentrallaboratoryIndividualheadsfromcommercialfish-eries sent to the central laboratoryhadbeenpreviously screenedelectronically for thepresenceofaCWTVoluntaryheadsamplesfromadiposefin-clippedindividualsfromrecreationalfisheriesweresenttothecentrallaboratorybutmanyoftheseindividualsdidnotcontainaCWTastherewasnopriorscreeningof these individu-als Stock composition of clipped individuals in the northern trollfishery has been previously outlined above Stock composition inthenorthern recreational fishery indicatedawidegeographicdis-tributionofcontributions rangingfromcoastalWashington (16)thewestcoastofVancouverIslandCU(10)theupperSkeenaCU(9)southeastAlaskaregion(8)mid-PugetSound(7)andeastcoast of Vancouver IslandndashStrait of Georgia CU (6) (SupportingInformation Table S4) Stock composition of clipped individuals inthecentralcoastrecreationalfisherywasdominatedbytheHoweSoundndashBurrardInletCU(27)andeastVancouverIslandndashStraitofGeorgiaCU(25)(SupportingInformationTableS4)butgiventhelowobservedclip rate in the fishery (014) therewasvery littlecatchthatcouldbeidentifiedashatcheryorigin
Directsamplingofrecreationalfisheries insouthernBCwhichincluded both clipped and unclipped individuals in the sample
Population Genotyped Assigned assignment
accuracy of population assignment
Robertson 66 66 1000 1000
Quinsam 69 68 986 1000
Puntledge 21 19 905 1000
BigQualicum 84 84 1000 1000
Goldstream 27 19 704 1000
Chilliwack 195 183 938 1000
Inch 91 85 934 1000
Stave 20 19 950 1000
Total 573 543 948 1000
TA B L E 3 emspPercentageofadiposefin-clippedjacks(age2years)sampledin2016andassignedviaparentageanalysistoparentsin2014broodstockpopulationsPopulationofsamplingwasassumedtobethepopulationoforiginofthejacksforevaluationofaccuracyofassignment
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp5BEACHAM Et Al
processed under the CWT recovery program for coho salmon inBCInadditionsampleswerecollectedfromarecreationalfishery(clippedandunclipped individuals) fromHaidaGwaiinearLangaraIslandarecreational fisherynearRivers Inlet (Area8clippedandunclippedindividuals)inthecentralcoastthepreviouslynotediceboat northern troll fishery and direct creel sampling (clipped andunclippedindividuals)ofrecreationalfisheriesinsouthernBC
Monthly catch composition estimates were determined forclipped individuals from various fisheries in BC Seasonal catchcomposition estimateswere determined by recording the numberofindividualgenotypesavailablefromthefisherysamplingineachmonth andweighting individualmonthly samples by the recordedcatch in themonth Individualswere randomlychosen formonthsinwhich the number of available genotypeswas in excess of thenumberrequiredfortheseasonalsampleForexampleif60oftheannualcatchwasrecordedinonemonthand200genotypeswereavailablefromthefisherythenallindividualssampledwouldbein-cludedintheseasonalsamplebutcontributionsofindividualsfromothermonthswerescaledrelativetothemonthlycontributiontothe
seasonalcatchIfthecatchinoneothermonthwas20oftheannualcatchand150sampleswereavailablethen200(2060)=67genotypeswererandomlychosenfromtheavailable150genotypesforinclusionintotheseasonalsample
InthenorthernBCtrollfisherysamplerselectronicallycheckedclippedindividualsforthepresenceofaCWTandheadsfromthoseindividualscontainingaCWTweresubsequentlysenttothecentrallaboratory for CWT recovery and tissue sampling for subsequentgenotypingTissuesamplesfromclippedindividualswithnoCWTsweredirectlyprovidedforgenotypingInordertoestimatemonthlyandseasonalstockcompositionofclippedindividualsinthisfisherywepooledthegenotypesobtainedfromdirectfisherysamplingofclippedindividualswiththosesubsequentlysenttothecentrallabo-ratoryforCWTrecoveryinordertoprovideanappropriatepoolofclippedindividualstoestimatemonthlyandseasonalstockcompo-sitionsofclippedindividualsinthefisheryIneachsampleanalyzedover all BC fisheries thenumberof individuals identified viaPBTrelative to thenumberofgenotypes in the samplewas tabulatedandsummarizedoversamplefisheryandseason
F I G U R E 1 emspMapindicatinggeographiclocationsforfisherysamplingand20populationsforwhichparentage-basedtaggingwasappliedinestimationofstockcomposition
6emsp |emsp emspensp BEACHAM Et Al
22emsp|emspEscapement sample collection
In2016adiposefin-clippedjacks(age2years)weresampledfromeightpopulationswith573individualscollectedandgenotypedsuc-cessfullyJacksweredefinedonthebasisofbodysizeduringsam-plecollectionwith jacksvisiblysmallerthanage3-yearspawnersPopulations sampled were as follows Robertson Creek QuinsamRiver Puntledge River Big Qualicum River Goldstream RiverChilliwackRiverStaveRiverand InchCreekTheobjectiveofthesamplingwas to obtain a sample of known origin and known agebaseduponknowncollectionlocationinordertoevaluateaccuracyofassignmentsof the jacks tobothCUandpopulationunder theassumption of no straying among populations for the individualssampled
In 2017 escapements (non-broodstock hatchery and river re-turns clipped individualsonly)were sampledwhere samplingwasfeasibleEscapementsamplingfrom13populationswasconductedand1692 individualsweregenotypedEscapement sampleswerenot available from all 20 populations forwhich itwas possible toassignindividualsinthenon-broodstockescapementsviaPBTTheobjectiveofthesamplingwastoevaluatestrayingratesamongpop-ulationsandforthosehatcheries(ChilliwackRiver)whereparentalgenotypeswereassignedtospecificreleasegroupsprovide infor-mationon the relative rates of returnof different release groupsAs relative ratesof return for the releasegroupswereconsideredancillarytothemainfocusofthestudynoresultsfromthesereturnswereincludedinthecurrentstudy
23emsp|emspExploitation rate
WeestimatedexploitationrateofadultcohosalmoninBCfisheriesviabothCWTsandgeneticsExploitationrateforapopulationwasdefinedasadultcatch(adultcatch-escapement)ForCWTstheob-servednumberofCWTswascorrectedbyldquono-pinrdquotaglossratesandwasexpandedbythepopulationrsquostag-specificmarkingratesummedovertagcodesandexpandedagainbythesamplingrateforthefish-eryinordertoestimatecatchofhatchery-originindividualsTheob-servednumberofCWTsinescapementsamplingwasexpandedinasimilarmanner inordertoestimatethehatcherycontributiontothe escapement For genetics the seasonal kept catchof adiposefin-clippedindividualsinafisherywasmultipliedbyaseasonalstockcompositionestimateinordertoestimatepopulation-specificcatchTheabundanceofhatchery-originescapementwascalculatedastheestimatedescapementmultipliedbytheproportionofadiposefin-clippedadultsobserved in theescapement If partof the juvenileproductionwasnotadiposefinclippeduponreleasefromthehatch-erythenthehatcherycontributionstobothcatchandescapementwereunderestimatedbythismethod
24emsp|emspGenetic stock identification baseline
The initial baseline was outlined by Beacham et al (2017) andconsisted of 20242 individuals from 117 populations with the
distributionofpopulationsrangingfromsoutheastAlaskatoPugetSoundinWashingtonStateThebaselinehasbeensubsequentlyex-panded to include 40774 individuals from267populations rang-ingfromsoutheastAlaskatoOregonTheprimaryexpansionofthebaselineincludedasurveyofadditionalpopulationsinsouthernBCcoastalWashingtontheColumbiaRiverdrainageandOregonThefull baseline is outlined in Supporting Information Table S1 withthe populations fromBC arrangedbyCU andwithUnited StatesofAmerica(US)populationsarrangedbygeographic(reporting)re-gionBeachametal (2017)hadpreviouslydemonstratedastrongregionalpopulationstructureandreportedthatoverall117popula-tionsaverageself-assignmentaccuracytopopulation(weightedbypopulation sample size)was 855 for individualswith an assign-mentprobabilitylt050excludedand928forindividualswithanassignmentprobabilitylt085excludedHighlevelsofaccuracyforGSI assignment of individuals to specific populations when com-binedwithassignmentsviaPBTprovideapowerful technique forassignmentofindividualsofunknownorigintosomepopulations
25emsp|emspLibrary preparation and genotyping
ThedetailedprocedureforlibrarypreparationandgenotypingwasoutlinedbyBeachametal(2017)SummarizedbrieflyDNAwasex-tractedfromcohosalmontissuesamplesviaaChelexextractionandtheDNAconcentration normalized to40ngμlwith a Tecan LiHarobotApanelofprimersdesignedtoamplifyspecificsegmentsofDNAthatcontainedSNPsofinterestwasappliedtotheextractedDNAwithprimersequencesforeachampliconoutlinedbyBeachametal (2017SupportingInformationTableS2)TheinitialmultiplexPCR amplification of 304 target ampliconswas conductedwith acocktail of 2μl of normalizedDNA extract 5μl of 2X Ion Agriseqprimerpool2μlofIonAgriseqHiFimixand1μlofddH2O Thermal cyclingwasconductedin96-wellPCRplates(oneindividualperwell)withthefollowingconditionsforPCR99degCndash2min17cycles(99degCndash15s60degCndash4min)10degChold
FollowingtheinitialPCRasecondstepemployingathermalcy-clerwasconducted thatpartiallydigested theprimerson theam-pliconsandthereactionconductedwiththefollowingconditions50degCndash10min55degCndash10min60degCndash20min10degCholdA thirdandfinalstepemployingathermalcyclerwasinitiatedtoligatethebarcodes(768individualcodes)totheampliconsandwasconductedwiththefollowingconditions22degCndash30min70degCndash10min10degCholdLibrarieswerepurifiedbyadditionof225μlofAgencourtregAMPureregXPmagneticbeadstoeachlibrarytheplatewasplacedonamagnetic racksupernatantdiscardedandthebeadswashedtwice in70ethanolThepurified librarieswerethenelutedwith25 μloflowTEand20μlofthesupernatanttransferredtoafresh96-well trayNext eachof the768prepared librarieswaspooledintoasingletubeforprocessingontheIonChefreg(ThermoFisherScientific)Twotubesofpooled librarieswereprocessedconsecu-tivelyontheIonChefandthus1536individualswereprocessedon a single run of the Ion ChefOne tube of the pooled librarieswasloadedontoeachP1regchipv3andthusampliconsfrom768
emspensp emsp | emsp7BEACHAM Et Al
individualsweredistributedoneachP1chipwith1536individualsprocessedbetweentwochipsThechipswerethenloadedontotheIonTorrentProtonsequencerAfter thesequencingrunwascom-pletedcomparisonswiththereferencegenomeoftherainbowtrout(O mykiss)(Berthelotetal2014)supplementedwiththesequencescontaining theobservedcohosalmonSNPswereconductedwithProton softwareVariant Callerreg and SNP genotypes at the sitesspecified by the hotspot filewithin target regionswere called byVariantCallerThehotspotfilecontained304SNPsiteswithoneSNPscoredateachampliconGenotypesatallavailableSNPsforanindividualwereassembled toprovideamulti-locus individualgen-otype The species identification SNPOkiOts_120255‐113 (StarksClementoampGarza2016)andsex identificationSNPOts_SEXY3‐1 wereomittedfromsubsequentparentageandGSIanalysesleaving302SNPs
26emsp|emspIdentification of individuals
AsnotedpreviouslyPBTandGSIwereusedconcurrently toesti-mate fishery stock composition Initially PBT was used and theanalysiswas conductedwhere the genotypes of individuals to beidentifiedwerematched to the genotypes of prospective parents(COLONY JonesampWang 2010Wang 2016) If all individuals ina hatchery broodstock are sampled and subsequently genotypedthenalloffspringfromthebroodstockaregeneticallymarkedThegenotypes of individuals of unknown origin are statistically com-pared with the genotypes of potential parents and if a match ismadetheoffspringareassignedtotheparentsandthustheori-ginandageoftheindividualsaredeterminedParentageassignmentsoftwarewasutilizedtoassignoffspringtoparentsasCOLONYcanproduceassignmentswhen thegenotypeofoneof theparents ismissingeitherduetoamissingparentalsampleor failure topro-duceaparentalgenotypefromanexistingsampleGiventhatPBTassignments for20potential populationswereevaluated for eachfisherysampleCOLONYwasrunwithallbroodstocksampleddur-ing2014inputasasingleunitforanalysisoffisherysampleswithnodifferentiationamongpopulationsAlthoughtheCOLONYassump-tionofasinglepopulationintheparentpoolwasviolatedanalysisofknown-originsamplesindicatedthatveryhighlevelsofaccuracywereachievedinassignmentswhenpoolingofpotentialparentsincontributing populationswas conducted Two-parent assignmentswere accepted only when both assigned parents originated fromthesamepopulationotherwisetheindividualwaspassedtogeneticstockidentification(GSI)forpotentialassignmentTwo-parentandsingle-parentassignmentswereacceptedonlywhentheprobabilityofcorrectassignmentwasge085fortheparentpairotherwisetheindividualwaspassedtopotentialassignmentbyGSIAdditionallyforsingle-parentassignments tobeaccepted thePBTassignmenttopopulationhad tobepart of theCUassigned to the individualviaGSI Individuals forwhichnoprospectiveparentswere identi-fiedinthebroodstockwerepassedtoGSIforpotentialassignmentPolygamousmatingwasassumedfortheCOLONYanalysisSimplepairwisecomparisonsbetweenoffspringandpotentialparentswere
conductedThebaselineforindividualssampledinthe2016escape-ments (jacks)and2017fisheries includedallbroodstockssampledin2014as these individualsarepredominately threeyearsofage(Sandercock1991)Jacks inthe2017escapementwere identifiedvia body size and subsequent assignment to parents in the 2015hatcherybroodstocksIndividualswithmorethan120missinggeno-typeswereeliminatedfromfurtheranalysesAnestimatedgenotyp-ingerrorrateof1wasusedforCOLONYassignmentsPreviouslyBeachametal(2017)hadreportedthatanaveragegenotypingerrorrate of 107 (1220discrepancies in 114105 comparisons) or analleleerrorrateof053(1220discrepanciesin228210compari-sons)wasobservedoverthe302SNPsscoredTheparentpairout-putfilewasthebasicfileusedinsubsequentanalyses
Thesecondmethodof individual identification isGSI inwhichthegeneticprofilesofwholepopulationspotentiallycontributingtoamixed-stocksampleareusedtoestimatetheoriginofeachindivid-ualinthesample(RUBIASMoranampAnderson2018)ThisanalysiswasrestrictedtothoseindividualsunassignedviaCOLONYForeachsample individuals not assigned by COLONYwere then assignedwithRUBIASThepopulationposteriormeansfilewasthebasicfileusedforsubsequentanalysesandthisfilecontainedtheprobabilityofassignmentoftheindividualtoeachofthe267populationsinthebaselineStockcompositionwasestimatedthroughthecombinationoffilesgeneratedwithbothCOLONYandRUBIAS Individualsas-signedviaCOLONYwereassignedaprobabilityof100oforiginat-ingfromtheidentifiedpopulationwitha000probabilityassignedtoallotherpopulationsinthebaselineThislevelofassignmentac-curacyviaPBTwasobservedpreviouslyforcohosalmonwiththepanelofSNPsemployedinthecurrentstudy(Beachametal2017)ThesedatawerethencombinedwiththeprobabilityofassignmentforthoseindividualsunassignedviaCOLONYtoeachofthepopu-lationsinthebaselineviaRUBIASAfteraninitialburn-inof25000iterations the last1000 iterations from theMonteCarloMarkovchainfromRUBIASwereusedtoestimatetheoriginofindividualsandstockcompositionwiththemeanallocationtoeachpopulationinthebaselinedeterminedStandarddeviationsofestimatedstockcompositionswere alsodetermined from the last1000 iterationsfrom theMonteCarloMarkovchain Stock compositionbyCUorreportinggroupwasdeterminedbysummationofallocationstoallpopulations in the baseline that belonged to the CU or reportinggroupunderconsideration
3emsp |emspRESULTS
31emsp|emspEstimation of stock composition for known‐origin samples
Genotypeswere available from573 jacks sampled in2016acrossthreeCUs and eight populations in southernBC Estimated stockcompositionbyCUand reportinggroup for this combinedsamplewas accuratewith an error utilizingonlyGSI of le04byCU forthreeCUspresentinthesample(Table2)WithbothPBTandGSIutilized theaverageerrordeclinedtole02byCUfor threeCUs
8emsp |emsp emspensp BEACHAM Et Al
TAB
LE 2
emspAccuracyofregional(UnitedStates)andconservationunit(Canada)estimatedstockcomposition()foramixed-stocksampleof573known-origincohosalmonjackssampledin
2016and620coded-wiretaggedindividualssampledin2017fisheriesinBCStandarddeviationoftheestimateisinparentheses
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
SoutheastAlaska
00
00(00)
00(00)
90
68(11)
68(11)
AlsekRiver
00
00(00)
00(00)
00
00(00)
00(00)
LowerStikine
00
00(00)
00(00)
00
04(03)
04(03)
LowerNass
00
00(00)
00(00)
63
64(10)
62(10)
UpperNass
00
00(00)
00(00)
00
04(03)
04(04)
PortlandSound-Observatory
Inlet-PortlandCanal
00
00(00)
00(00)
00
00(00)
00(00)
SkeenaEstuary
00
00(00)
00(00)
00
00(00)
00(00)
LowerSkeena
00
00(00)
00(00)
16
12(05)
12(05)
MiddleSkeena
00
00(00)
00(00)
69
39(08)
39(09)
UpperSkeena
00
00(00)
00(00)
06
40(10)
40(09)
HaidaGwaiindashGrahamIslandLowlands
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-East
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-W
est
00
00(00)
00(00)
00
00(00)
00(00)
NorthernCoastalStreams
00
00(00)
00(00)
00
00(00)
00(01)
HecateStraitMainland
00
00(00)
00(00)
00
07(05)
06(04)
MusselndashKynoch
00
00(00)
00(00)
00
00(00)
00(00)
DouglasChannelndashKitimatArm
00
00(01)
00(01)
00
00(01)
00(01)
BellaCoolandashDeanRivers
00
00(00)
00(00)
00
00(00)
00(00)
RiversInlet
00
00(01)
00(00)
00
00(00)
00(00)
SmithInlet
00
00(00)
00(00)
00
00(00)
00(00)
SouthernCoastalStreamsndashQueen
CharlotteStraitndashJohnstoneStraitndash
SouthernFjords
00
00(01)
00(01)
00
00(02)
00(02)
HomathkondashKlinakliniRivers
00
00(00)
00(00)
00
00(00)
00(00)
GeorgiaStraitMainland
00
00(00)
00(00)
00
00(00)
00(00)
HoweSoundndashBurrardInlet
00
00(00)
00(00)
06
00(01)
00(00)
EastVancouverIslandndashGeorgiaStrait
351
345(20)
348(20)
131
134(14)
140(14)
EastVancouverIslandndashJohnstone
StraitndashSouthernFjords
00
00(00)
00(00)
00
00(01)
00(00)
NahwittiLowland
00
00(01)
00(00)
18
17(06)
17(06) (C
ontin
ues)
emspensp emsp | emsp9BEACHAM Et Al
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
WestVancouverIsland
115
115(14)
115(13)
135
134(14)
135(14)
Clayoquot
00
00(00)
00(00)
00
02(03)
02(02)
JuandeFucandashPachena
00
00(01)
00(01)
00
00(00)
00(00)
LowerFraser
534
539(21)
536(21)
185
187(15)
188(15)
Lillooet
00
00(00)
00(00)
00
03(02)
02(01)
FraserCanyon
00
00(00)
00(00)
00
00(00)
00(00)
InteriorFraser
00
00(00)
00(00)
00
00(00)
00(00)
LowerThompson
00
00(00)
00(00)
08
07(03)
08(04)
NorthThompson
00
00(00)
00(01)
00
00(00)
00(00)
SouthThompson
00
00(00)
00(00)
05
06(03)
05(03)
Boun
dary
Bay
00
00(00)
00(00)
00
00(00)
00(00)
NooksackRiver
00
01(02)
01(02)
26
09(09)
10(08)
SkagitRiver
00
00(01)
00(01)
05
23(08)
22(08)
NorthernPugetSound
00
00(00)
00(00)
39
58(12)
55(13)
Mid-PugetSound
00
00(00)
00(00)
52
62(11)
63(11)
SouthernPugetSound
00
00(00)
00(00)
10
09(10)
06(08)
JuandeFucaStrait
00
00(00)
00(00)
00
02(03)
01(03)
HoodCanal
00
00(00)
00(00)
44
16(05)
16(06)
CoastalWashington
00
00(00)
00(00)
60
59(10)
59(10)
ColumbiaRiver
00
00(00)
00(00)
32
32(07)
32(07)
Oregon
00
00(00)
00(00)
00
00(01)
00(00)
TAB
LE 2
(Continued)
10emsp |emsp emspensp BEACHAM Et Al
presentinthesample(Table2)Theaverageerrorfor45CUsorre-portinggroupsabsentinthesamplewas00bybothGSIandPBT-GSIBypopulation543(948)ofthejackswereassignedviaPBTwith 100 accuracywith respect to population of origin and age(Table3)Estimatedstockcompositionfor20populationswhereitwaspossibletousebothPBTandGSIinestimationofstockcompo-sitionwasaccurateFortheeightpopulationspresentinthesampleofjackstheaverageerrorutilizingGSIonlywas05perpopulationdecliningto02utilizingbothPBTandGSIundertheassumptionthatthecollectionlocationwasanaccuratereflectionofjackoriginTherewasanaverage00errorforthe12populationswithnorep-resentation inthesamplewithutilizationofeitherGSIorPBT-GSI(Table4)ThelargesterrorutilizingbothPBTandGSI(underestima-tionby06ofactualvalue)wasobservedintheGoldstreamRiverpopulationwhichalsodisplayedthe lowestpercentage (704)ofjacksidentifiedviaPBT(Table3)PBTwhencombinedwithGSIpro-ducedmoreaccurateestimatesofCU-specificandpopulation-spe-cificstockcompositionthanavailablewithonlyGSI
Of the 3054 adipose fin-clipped coho salmon samples re-ceivedforpotentialgenotypingfromacentrallaboratorywhere2017fisherysampleswereprocessedforpotentialCWTrecov-erygenotypeswereobtained for2533 (829)of the individ-uals processed There were 750 CWTs recovered from 3054headsor snouts examined and genotypeswereobtained from622 (829) of the individuals that carried a CWT GenotypesofcohosalmonwithoutCWTsbutsentfromtheCWTheadre-covery laboratorywereobtainedfrom1911of2304 individu-als (829) Release information associatedwith theCWTwasavailable for 620 of the 622CWTs associatedwith genotypedindividuals and these 620 individuals constituted a sample ofknown origin There were individuals from 20 regions or CUspresentinthissampleandforthese20regionsorCUstheav-erageerrorofestimationutilizingonlyGSIwas09perregionorCUand09forbothPBTandGSIcombined(Table2)CWTswererecoveredoriginatingfromsevenpopulationswhereitwaspossibletoincorporatebothGSIandPBTinestimationofstockcompositionFor these sevenpopulations theaverageerror inestimatedstockcompositionsperpopulationwas20withGSIand 03 with PBT-GSI (Table 4) There was an average 00errorforthe13populationswithnorepresentationinthesam-ple with utilization of either GSI or PBT-GSI (Table 4) Similartothesampleof jacksPBTwhencombinedwithGSIproducedmore accurate estimates of population-specific stock composi-tionthanavailablewithonlyGSI
32emsp|emspEvaluation of 2016 fishery sampling
Asbroodstocksamplingatselectedhatcheries insouthernBCdidnot commence until 2014 it was not possible to obtain PBT as-signments for any fishery samples collected in 2016 HoweverGSI analysis of individuals derived from recreational fishery sam-ples intheStraitofGeorgiaJuandeFucaStraitandoffthewestcoastofVancouverIslandinsouthernBCindicatedthattherewas
virtually no contribution from regions orCUsnorthofVancouverIsland (Supporting Information Table S2) The only exceptionwasin thenorthwestcoastofVancouver Islandboth inshoreandoff-shorewhereupto12ofthesamplewasidentifiedasoriginatingfromnorthernandcentralBCCUsUS-origincohosalmonwerees-timatedtohavecontributedapproximately70ofsummerrecrea-tionalsamplesderivedfromoffshorefisheriesalongthewestcoastofVancouverIslandaswellasintheentrancetoJuandeFucaStrait(A21A121)InmoreinshoreareasalongthewestcoastofVancouverIslandcohosalmonfromlocalCUswerethedominantcontributorstothefisheryForexamplethewestcoastofVancouverIslandCUcontributedapproximately80ofthe39-individualAugustsamplefromtheinshoresouthwesternVancouverIslandregionIntheStraitofGeorgia localCUswereagainthemaincontributorstotherec-reationalfisheryduringthesummerwiththeHoweSoundndashBurrardInletCUcontributingupto40insomesamplestheeastcoastofVancouverIslandCUuptoapproximately30andthelowerFraserRiverCUuptoapproximately40WhilethelowerFraserRiverCUwasanimportantcontributortotherecreationalfisheryintheStraitofGeorgiaandJuandeFucaStraititscontributiontofisheriesoffthewestcoastofVancouverIslandwaslimitedTheNooksackRiverregion innorthernWashingtonStatewasalso identifiedasacon-sistent contributor to recreational fishery samples in the Strait ofGeorgia insomecasescontributingupto20ofthesampleana-lyzed(SupportingInformationTableS2)
StockcompositionswereavailablefromtwotestfisheriesoneatrolltestfisherynearBrooksPeninsulabynorthwesternVancouverIslandandtheotheragillnettestfisheryinupperJohnstoneStraitIn the Brooks Peninsula fishery Vancouver IslandCUs comprised40ndash60ofthesampleswithaminorcontributionofFraserRiverCUs (lt10) The five interior Fraser River CUs of conservationconcern comprised00ndash33of the sampleswhereas themoreabundant lower Fraser River CUs comprised only 00ndash55 ofthesamples Individuals fromFraserRiverCUswerenotdetectedduringJulyinArea127theoffshoreportionofnorthernnorthwest-ernVancouverIslandbuttheyweredetectedinArea126themoresoutherlyoffshoreportionofnorthwesternVancouverIslandIntheJohnstoneStrait(RoundIsland)testfisheryindividualsfromFraserRiverCUswerescarcebutthefiveinteriorFraserRiverCUscom-prised30ndash35of the sampleswhereas the lowerFraserRiverCUscomprisedonly00ndash65ofthesamplesThemajorcontrib-utors to the samplesoriginated from theeast coastofVancouverIsland(24ndash45)andHoweSoundndashBurrardInletCUs(9ndash11)aswellascoastalWashington(8ndash15)(SupportingInformationTableS2)
ThefinalfisherysampledwasademonstrationcommercialtrollfisheryincentralcoastalBC(A6andA7)ContributionsfromlocalCUswerehigherinsamplesfrommoreinshoreareas(A7-7August212southernCUsandregions)withcontributionsfromsouth-ernCUshigherinthemoreseawardareas(A6-9September862southernCUsandregions)LikethewestcoastofVancouverIslandmigrating stocks were more likely to be found in more seawardlocations
emspensp emsp | emsp11BEACHAM Et Al
33emsp|emspEvaluation of 2017 fishery sampling
Onesourceoffisherysampleswasdirectfisherysamplingofindi-viduals(bothadiposefin-clippedandunclipped)wherethefocuswas not associated with potential CWT recovery In this casesamplingofunclippedindividualsfromthenorthern(AreaF)trollfishery indicated that initially (July 10ndash12) northern and centralBCCUscontributed23tothecatchsouthernBCCUs53andsouthern US regions 23 (Supporting Information Table S3)Stock composition changed markedly in samples collected July13ndash18 with Alaska contributing 12 northern and central BCCUs contributing 72 and southern BCCUs contributing 16withnosouthernUScontributionSamplescollectedduringJuly22ndash28revealedadominantcontributionfromnorthernandcen-tralBCCUs(96)andCUsfromthisregioncontributedbyfarthelargestportionof the catch in subsequent sampleswith south-ernBCCUsandsouthernUSregionslargelyabsent(SupportingInformation Table S3) The average contribution to the weeklysamples from July 22 onwardswas 17 southeast Alaska 16HaidaGwaiindashGrahamIslandLowlandsCU15DouglasChannelndashKitimatArmCU 12Hecate StraitMainlandCU 8NorthernCoastalStreamsCUand7LowerNassCUNotunexpectedlysampling of only clipped individuals from the fishery indicatedverysubstantialdifferencesinstockcompositionbetweenclippedandunclippedindividualsTheaveragecontributiontotheweeklysamples of clipped individuals from July 19 onwards was 32coastalWashington11mid-PugetSound10westVancouverIsland CU and 5 for each of northern Puget Sound and Juande Fuca Strait (Supporting Information Table S4) However theobservedcliprateinthefisherywas16sothecatchwasdomi-natedbyunclippedindividualsTheSeptember2ndash8samplefromthe Haida Gwaii recreational fishery near Langara displayed astockcompositionof42HaidaGwaiindashGrahamIslandLowlandsCU15HecateStraitMainlandCU10LowerSkeenaCUand8 Northern Coastal Streams CU similar to that observed inthe troll fisherywith the local CU (HaidaGwaiindashGraham IslandLowlands) inhigherproportionsaswouldbeexpectedinamoreinshorefishery(SupportingInformationTableS3)
Directsamplingof thecentralcoast troll fisheryrevealedthatonaverage theDouglasChannelndashKitimatArmCUwasthe largestcontributortothefishery(24)followedbytheNorthernCoastalStreamsCU(19)BellaCoolandashDeanRivers(18)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(10) (Supporting InformationTableS3)withallCUsproximaltothelocationofthefisherySamplesfromtherecreationalcentralcoastfisheryindicatedthattheBellaCoolandashDeanRiversCUwasthedominant contributor to the fishery (32) followedby theRiversInletCU(19)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(15)andDouglasChannelndashKitimatArmCU(7) (SupportingInformationTableS3)GiventhelocationofthefisherydominantcontributionsfromtheseCUsweretobeexpected
AsnotedpreviouslysampleswerealsoavailablefromvirtuallyallindividualssampledfromcommercialrecreationalandFirstNationsfisheriesthatwouldberoutinelyprocessedforCWTrecoveryinayearbyacentrallaboratoryIndividualheadsfromcommercialfish-eries sent to the central laboratoryhadbeenpreviously screenedelectronically for thepresenceofaCWTVoluntaryheadsamplesfromadiposefin-clippedindividualsfromrecreationalfisheriesweresenttothecentrallaboratorybutmanyoftheseindividualsdidnotcontainaCWTastherewasnopriorscreeningof these individu-als Stock composition of clipped individuals in the northern trollfishery has been previously outlined above Stock composition inthenorthern recreational fishery indicatedawidegeographicdis-tributionofcontributions rangingfromcoastalWashington (16)thewestcoastofVancouverIslandCU(10)theupperSkeenaCU(9)southeastAlaskaregion(8)mid-PugetSound(7)andeastcoast of Vancouver IslandndashStrait of Georgia CU (6) (SupportingInformation Table S4) Stock composition of clipped individuals inthecentralcoastrecreationalfisherywasdominatedbytheHoweSoundndashBurrardInletCU(27)andeastVancouverIslandndashStraitofGeorgiaCU(25)(SupportingInformationTableS4)butgiventhelowobservedclip rate in the fishery (014) therewasvery littlecatchthatcouldbeidentifiedashatcheryorigin
Directsamplingofrecreationalfisheries insouthernBCwhichincluded both clipped and unclipped individuals in the sample
Population Genotyped Assigned assignment
accuracy of population assignment
Robertson 66 66 1000 1000
Quinsam 69 68 986 1000
Puntledge 21 19 905 1000
BigQualicum 84 84 1000 1000
Goldstream 27 19 704 1000
Chilliwack 195 183 938 1000
Inch 91 85 934 1000
Stave 20 19 950 1000
Total 573 543 948 1000
TA B L E 3 emspPercentageofadiposefin-clippedjacks(age2years)sampledin2016andassignedviaparentageanalysistoparentsin2014broodstockpopulationsPopulationofsamplingwasassumedtobethepopulationoforiginofthejacksforevaluationofaccuracyofassignment
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
6emsp |emsp emspensp BEACHAM Et Al
22emsp|emspEscapement sample collection
In2016adiposefin-clippedjacks(age2years)weresampledfromeightpopulationswith573individualscollectedandgenotypedsuc-cessfullyJacksweredefinedonthebasisofbodysizeduringsam-plecollectionwith jacksvisiblysmallerthanage3-yearspawnersPopulations sampled were as follows Robertson Creek QuinsamRiver Puntledge River Big Qualicum River Goldstream RiverChilliwackRiverStaveRiverand InchCreekTheobjectiveofthesamplingwas to obtain a sample of known origin and known agebaseduponknowncollectionlocationinordertoevaluateaccuracyofassignmentsof the jacks tobothCUandpopulationunder theassumption of no straying among populations for the individualssampled
In 2017 escapements (non-broodstock hatchery and river re-turns clipped individualsonly)were sampledwhere samplingwasfeasibleEscapementsamplingfrom13populationswasconductedand1692 individualsweregenotypedEscapement sampleswerenot available from all 20 populations forwhich itwas possible toassignindividualsinthenon-broodstockescapementsviaPBTTheobjectiveofthesamplingwastoevaluatestrayingratesamongpop-ulationsandforthosehatcheries(ChilliwackRiver)whereparentalgenotypeswereassignedtospecificreleasegroupsprovide infor-mationon the relative rates of returnof different release groupsAs relative ratesof return for the releasegroupswereconsideredancillarytothemainfocusofthestudynoresultsfromthesereturnswereincludedinthecurrentstudy
23emsp|emspExploitation rate
WeestimatedexploitationrateofadultcohosalmoninBCfisheriesviabothCWTsandgeneticsExploitationrateforapopulationwasdefinedasadultcatch(adultcatch-escapement)ForCWTstheob-servednumberofCWTswascorrectedbyldquono-pinrdquotaglossratesandwasexpandedbythepopulationrsquostag-specificmarkingratesummedovertagcodesandexpandedagainbythesamplingrateforthefish-eryinordertoestimatecatchofhatchery-originindividualsTheob-servednumberofCWTsinescapementsamplingwasexpandedinasimilarmanner inordertoestimatethehatcherycontributiontothe escapement For genetics the seasonal kept catchof adiposefin-clippedindividualsinafisherywasmultipliedbyaseasonalstockcompositionestimateinordertoestimatepopulation-specificcatchTheabundanceofhatchery-originescapementwascalculatedastheestimatedescapementmultipliedbytheproportionofadiposefin-clippedadultsobserved in theescapement If partof the juvenileproductionwasnotadiposefinclippeduponreleasefromthehatch-erythenthehatcherycontributionstobothcatchandescapementwereunderestimatedbythismethod
24emsp|emspGenetic stock identification baseline
The initial baseline was outlined by Beacham et al (2017) andconsisted of 20242 individuals from 117 populations with the
distributionofpopulationsrangingfromsoutheastAlaskatoPugetSoundinWashingtonStateThebaselinehasbeensubsequentlyex-panded to include 40774 individuals from267populations rang-ingfromsoutheastAlaskatoOregonTheprimaryexpansionofthebaselineincludedasurveyofadditionalpopulationsinsouthernBCcoastalWashingtontheColumbiaRiverdrainageandOregonThefull baseline is outlined in Supporting Information Table S1 withthe populations fromBC arrangedbyCU andwithUnited StatesofAmerica(US)populationsarrangedbygeographic(reporting)re-gionBeachametal (2017)hadpreviouslydemonstratedastrongregionalpopulationstructureandreportedthatoverall117popula-tionsaverageself-assignmentaccuracytopopulation(weightedbypopulation sample size)was 855 for individualswith an assign-mentprobabilitylt050excludedand928forindividualswithanassignmentprobabilitylt085excludedHighlevelsofaccuracyforGSI assignment of individuals to specific populations when com-binedwithassignmentsviaPBTprovideapowerful technique forassignmentofindividualsofunknownorigintosomepopulations
25emsp|emspLibrary preparation and genotyping
ThedetailedprocedureforlibrarypreparationandgenotypingwasoutlinedbyBeachametal(2017)SummarizedbrieflyDNAwasex-tractedfromcohosalmontissuesamplesviaaChelexextractionandtheDNAconcentration normalized to40ngμlwith a Tecan LiHarobotApanelofprimersdesignedtoamplifyspecificsegmentsofDNAthatcontainedSNPsofinterestwasappliedtotheextractedDNAwithprimersequencesforeachampliconoutlinedbyBeachametal (2017SupportingInformationTableS2)TheinitialmultiplexPCR amplification of 304 target ampliconswas conductedwith acocktail of 2μl of normalizedDNA extract 5μl of 2X Ion Agriseqprimerpool2μlofIonAgriseqHiFimixand1μlofddH2O Thermal cyclingwasconductedin96-wellPCRplates(oneindividualperwell)withthefollowingconditionsforPCR99degCndash2min17cycles(99degCndash15s60degCndash4min)10degChold
FollowingtheinitialPCRasecondstepemployingathermalcy-clerwasconducted thatpartiallydigested theprimerson theam-pliconsandthereactionconductedwiththefollowingconditions50degCndash10min55degCndash10min60degCndash20min10degCholdA thirdandfinalstepemployingathermalcyclerwasinitiatedtoligatethebarcodes(768individualcodes)totheampliconsandwasconductedwiththefollowingconditions22degCndash30min70degCndash10min10degCholdLibrarieswerepurifiedbyadditionof225μlofAgencourtregAMPureregXPmagneticbeadstoeachlibrarytheplatewasplacedonamagnetic racksupernatantdiscardedandthebeadswashedtwice in70ethanolThepurified librarieswerethenelutedwith25 μloflowTEand20μlofthesupernatanttransferredtoafresh96-well trayNext eachof the768prepared librarieswaspooledintoasingletubeforprocessingontheIonChefreg(ThermoFisherScientific)Twotubesofpooled librarieswereprocessedconsecu-tivelyontheIonChefandthus1536individualswereprocessedon a single run of the Ion ChefOne tube of the pooled librarieswasloadedontoeachP1regchipv3andthusampliconsfrom768
emspensp emsp | emsp7BEACHAM Et Al
individualsweredistributedoneachP1chipwith1536individualsprocessedbetweentwochipsThechipswerethenloadedontotheIonTorrentProtonsequencerAfter thesequencingrunwascom-pletedcomparisonswiththereferencegenomeoftherainbowtrout(O mykiss)(Berthelotetal2014)supplementedwiththesequencescontaining theobservedcohosalmonSNPswereconductedwithProton softwareVariant Callerreg and SNP genotypes at the sitesspecified by the hotspot filewithin target regionswere called byVariantCallerThehotspotfilecontained304SNPsiteswithoneSNPscoredateachampliconGenotypesatallavailableSNPsforanindividualwereassembled toprovideamulti-locus individualgen-otype The species identification SNPOkiOts_120255‐113 (StarksClementoampGarza2016)andsex identificationSNPOts_SEXY3‐1 wereomittedfromsubsequentparentageandGSIanalysesleaving302SNPs
26emsp|emspIdentification of individuals
AsnotedpreviouslyPBTandGSIwereusedconcurrently toesti-mate fishery stock composition Initially PBT was used and theanalysiswas conductedwhere the genotypes of individuals to beidentifiedwerematched to the genotypes of prospective parents(COLONY JonesampWang 2010Wang 2016) If all individuals ina hatchery broodstock are sampled and subsequently genotypedthenalloffspringfromthebroodstockaregeneticallymarkedThegenotypes of individuals of unknown origin are statistically com-pared with the genotypes of potential parents and if a match ismadetheoffspringareassignedtotheparentsandthustheori-ginandageoftheindividualsaredeterminedParentageassignmentsoftwarewasutilizedtoassignoffspringtoparentsasCOLONYcanproduceassignmentswhen thegenotypeofoneof theparents ismissingeitherduetoamissingparentalsampleor failure topro-duceaparentalgenotypefromanexistingsampleGiventhatPBTassignments for20potential populationswereevaluated for eachfisherysampleCOLONYwasrunwithallbroodstocksampleddur-ing2014inputasasingleunitforanalysisoffisherysampleswithnodifferentiationamongpopulationsAlthoughtheCOLONYassump-tionofasinglepopulationintheparentpoolwasviolatedanalysisofknown-originsamplesindicatedthatveryhighlevelsofaccuracywereachievedinassignmentswhenpoolingofpotentialparentsincontributing populationswas conducted Two-parent assignmentswere accepted only when both assigned parents originated fromthesamepopulationotherwisetheindividualwaspassedtogeneticstockidentification(GSI)forpotentialassignmentTwo-parentandsingle-parentassignmentswereacceptedonlywhentheprobabilityofcorrectassignmentwasge085fortheparentpairotherwisetheindividualwaspassedtopotentialassignmentbyGSIAdditionallyforsingle-parentassignments tobeaccepted thePBTassignmenttopopulationhad tobepart of theCUassigned to the individualviaGSI Individuals forwhichnoprospectiveparentswere identi-fiedinthebroodstockwerepassedtoGSIforpotentialassignmentPolygamousmatingwasassumedfortheCOLONYanalysisSimplepairwisecomparisonsbetweenoffspringandpotentialparentswere
conductedThebaselineforindividualssampledinthe2016escape-ments (jacks)and2017fisheries includedallbroodstockssampledin2014as these individualsarepredominately threeyearsofage(Sandercock1991)Jacks inthe2017escapementwere identifiedvia body size and subsequent assignment to parents in the 2015hatcherybroodstocksIndividualswithmorethan120missinggeno-typeswereeliminatedfromfurtheranalysesAnestimatedgenotyp-ingerrorrateof1wasusedforCOLONYassignmentsPreviouslyBeachametal(2017)hadreportedthatanaveragegenotypingerrorrate of 107 (1220discrepancies in 114105 comparisons) or analleleerrorrateof053(1220discrepanciesin228210compari-sons)wasobservedoverthe302SNPsscoredTheparentpairout-putfilewasthebasicfileusedinsubsequentanalyses
Thesecondmethodof individual identification isGSI inwhichthegeneticprofilesofwholepopulationspotentiallycontributingtoamixed-stocksampleareusedtoestimatetheoriginofeachindivid-ualinthesample(RUBIASMoranampAnderson2018)ThisanalysiswasrestrictedtothoseindividualsunassignedviaCOLONYForeachsample individuals not assigned by COLONYwere then assignedwithRUBIASThepopulationposteriormeansfilewasthebasicfileusedforsubsequentanalysesandthisfilecontainedtheprobabilityofassignmentoftheindividualtoeachofthe267populationsinthebaselineStockcompositionwasestimatedthroughthecombinationoffilesgeneratedwithbothCOLONYandRUBIAS Individualsas-signedviaCOLONYwereassignedaprobabilityof100oforiginat-ingfromtheidentifiedpopulationwitha000probabilityassignedtoallotherpopulationsinthebaselineThislevelofassignmentac-curacyviaPBTwasobservedpreviouslyforcohosalmonwiththepanelofSNPsemployedinthecurrentstudy(Beachametal2017)ThesedatawerethencombinedwiththeprobabilityofassignmentforthoseindividualsunassignedviaCOLONYtoeachofthepopu-lationsinthebaselineviaRUBIASAfteraninitialburn-inof25000iterations the last1000 iterations from theMonteCarloMarkovchainfromRUBIASwereusedtoestimatetheoriginofindividualsandstockcompositionwiththemeanallocationtoeachpopulationinthebaselinedeterminedStandarddeviationsofestimatedstockcompositionswere alsodetermined from the last1000 iterationsfrom theMonteCarloMarkovchain Stock compositionbyCUorreportinggroupwasdeterminedbysummationofallocationstoallpopulations in the baseline that belonged to the CU or reportinggroupunderconsideration
3emsp |emspRESULTS
31emsp|emspEstimation of stock composition for known‐origin samples
Genotypeswere available from573 jacks sampled in2016acrossthreeCUs and eight populations in southernBC Estimated stockcompositionbyCUand reportinggroup for this combinedsamplewas accuratewith an error utilizingonlyGSI of le04byCU forthreeCUspresentinthesample(Table2)WithbothPBTandGSIutilized theaverageerrordeclinedtole02byCUfor threeCUs
8emsp |emsp emspensp BEACHAM Et Al
TAB
LE 2
emspAccuracyofregional(UnitedStates)andconservationunit(Canada)estimatedstockcomposition()foramixed-stocksampleof573known-origincohosalmonjackssampledin
2016and620coded-wiretaggedindividualssampledin2017fisheriesinBCStandarddeviationoftheestimateisinparentheses
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
SoutheastAlaska
00
00(00)
00(00)
90
68(11)
68(11)
AlsekRiver
00
00(00)
00(00)
00
00(00)
00(00)
LowerStikine
00
00(00)
00(00)
00
04(03)
04(03)
LowerNass
00
00(00)
00(00)
63
64(10)
62(10)
UpperNass
00
00(00)
00(00)
00
04(03)
04(04)
PortlandSound-Observatory
Inlet-PortlandCanal
00
00(00)
00(00)
00
00(00)
00(00)
SkeenaEstuary
00
00(00)
00(00)
00
00(00)
00(00)
LowerSkeena
00
00(00)
00(00)
16
12(05)
12(05)
MiddleSkeena
00
00(00)
00(00)
69
39(08)
39(09)
UpperSkeena
00
00(00)
00(00)
06
40(10)
40(09)
HaidaGwaiindashGrahamIslandLowlands
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-East
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-W
est
00
00(00)
00(00)
00
00(00)
00(00)
NorthernCoastalStreams
00
00(00)
00(00)
00
00(00)
00(01)
HecateStraitMainland
00
00(00)
00(00)
00
07(05)
06(04)
MusselndashKynoch
00
00(00)
00(00)
00
00(00)
00(00)
DouglasChannelndashKitimatArm
00
00(01)
00(01)
00
00(01)
00(01)
BellaCoolandashDeanRivers
00
00(00)
00(00)
00
00(00)
00(00)
RiversInlet
00
00(01)
00(00)
00
00(00)
00(00)
SmithInlet
00
00(00)
00(00)
00
00(00)
00(00)
SouthernCoastalStreamsndashQueen
CharlotteStraitndashJohnstoneStraitndash
SouthernFjords
00
00(01)
00(01)
00
00(02)
00(02)
HomathkondashKlinakliniRivers
00
00(00)
00(00)
00
00(00)
00(00)
GeorgiaStraitMainland
00
00(00)
00(00)
00
00(00)
00(00)
HoweSoundndashBurrardInlet
00
00(00)
00(00)
06
00(01)
00(00)
EastVancouverIslandndashGeorgiaStrait
351
345(20)
348(20)
131
134(14)
140(14)
EastVancouverIslandndashJohnstone
StraitndashSouthernFjords
00
00(00)
00(00)
00
00(01)
00(00)
NahwittiLowland
00
00(01)
00(00)
18
17(06)
17(06) (C
ontin
ues)
emspensp emsp | emsp9BEACHAM Et Al
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
WestVancouverIsland
115
115(14)
115(13)
135
134(14)
135(14)
Clayoquot
00
00(00)
00(00)
00
02(03)
02(02)
JuandeFucandashPachena
00
00(01)
00(01)
00
00(00)
00(00)
LowerFraser
534
539(21)
536(21)
185
187(15)
188(15)
Lillooet
00
00(00)
00(00)
00
03(02)
02(01)
FraserCanyon
00
00(00)
00(00)
00
00(00)
00(00)
InteriorFraser
00
00(00)
00(00)
00
00(00)
00(00)
LowerThompson
00
00(00)
00(00)
08
07(03)
08(04)
NorthThompson
00
00(00)
00(01)
00
00(00)
00(00)
SouthThompson
00
00(00)
00(00)
05
06(03)
05(03)
Boun
dary
Bay
00
00(00)
00(00)
00
00(00)
00(00)
NooksackRiver
00
01(02)
01(02)
26
09(09)
10(08)
SkagitRiver
00
00(01)
00(01)
05
23(08)
22(08)
NorthernPugetSound
00
00(00)
00(00)
39
58(12)
55(13)
Mid-PugetSound
00
00(00)
00(00)
52
62(11)
63(11)
SouthernPugetSound
00
00(00)
00(00)
10
09(10)
06(08)
JuandeFucaStrait
00
00(00)
00(00)
00
02(03)
01(03)
HoodCanal
00
00(00)
00(00)
44
16(05)
16(06)
CoastalWashington
00
00(00)
00(00)
60
59(10)
59(10)
ColumbiaRiver
00
00(00)
00(00)
32
32(07)
32(07)
Oregon
00
00(00)
00(00)
00
00(01)
00(00)
TAB
LE 2
(Continued)
10emsp |emsp emspensp BEACHAM Et Al
presentinthesample(Table2)Theaverageerrorfor45CUsorre-portinggroupsabsentinthesamplewas00bybothGSIandPBT-GSIBypopulation543(948)ofthejackswereassignedviaPBTwith 100 accuracywith respect to population of origin and age(Table3)Estimatedstockcompositionfor20populationswhereitwaspossibletousebothPBTandGSIinestimationofstockcompo-sitionwasaccurateFortheeightpopulationspresentinthesampleofjackstheaverageerrorutilizingGSIonlywas05perpopulationdecliningto02utilizingbothPBTandGSIundertheassumptionthatthecollectionlocationwasanaccuratereflectionofjackoriginTherewasanaverage00errorforthe12populationswithnorep-resentation inthesamplewithutilizationofeitherGSIorPBT-GSI(Table4)ThelargesterrorutilizingbothPBTandGSI(underestima-tionby06ofactualvalue)wasobservedintheGoldstreamRiverpopulationwhichalsodisplayedthe lowestpercentage (704)ofjacksidentifiedviaPBT(Table3)PBTwhencombinedwithGSIpro-ducedmoreaccurateestimatesofCU-specificandpopulation-spe-cificstockcompositionthanavailablewithonlyGSI
Of the 3054 adipose fin-clipped coho salmon samples re-ceivedforpotentialgenotypingfromacentrallaboratorywhere2017fisherysampleswereprocessedforpotentialCWTrecov-erygenotypeswereobtained for2533 (829)of the individ-uals processed There were 750 CWTs recovered from 3054headsor snouts examined and genotypeswereobtained from622 (829) of the individuals that carried a CWT GenotypesofcohosalmonwithoutCWTsbutsentfromtheCWTheadre-covery laboratorywereobtainedfrom1911of2304 individu-als (829) Release information associatedwith theCWTwasavailable for 620 of the 622CWTs associatedwith genotypedindividuals and these 620 individuals constituted a sample ofknown origin There were individuals from 20 regions or CUspresentinthissampleandforthese20regionsorCUstheav-erageerrorofestimationutilizingonlyGSIwas09perregionorCUand09forbothPBTandGSIcombined(Table2)CWTswererecoveredoriginatingfromsevenpopulationswhereitwaspossibletoincorporatebothGSIandPBTinestimationofstockcompositionFor these sevenpopulations theaverageerror inestimatedstockcompositionsperpopulationwas20withGSIand 03 with PBT-GSI (Table 4) There was an average 00errorforthe13populationswithnorepresentationinthesam-ple with utilization of either GSI or PBT-GSI (Table 4) Similartothesampleof jacksPBTwhencombinedwithGSIproducedmore accurate estimates of population-specific stock composi-tionthanavailablewithonlyGSI
32emsp|emspEvaluation of 2016 fishery sampling
Asbroodstocksamplingatselectedhatcheries insouthernBCdidnot commence until 2014 it was not possible to obtain PBT as-signments for any fishery samples collected in 2016 HoweverGSI analysis of individuals derived from recreational fishery sam-ples intheStraitofGeorgiaJuandeFucaStraitandoffthewestcoastofVancouverIslandinsouthernBCindicatedthattherewas
virtually no contribution from regions orCUsnorthofVancouverIsland (Supporting Information Table S2) The only exceptionwasin thenorthwestcoastofVancouver Islandboth inshoreandoff-shorewhereupto12ofthesamplewasidentifiedasoriginatingfromnorthernandcentralBCCUsUS-origincohosalmonwerees-timatedtohavecontributedapproximately70ofsummerrecrea-tionalsamplesderivedfromoffshorefisheriesalongthewestcoastofVancouverIslandaswellasintheentrancetoJuandeFucaStrait(A21A121)InmoreinshoreareasalongthewestcoastofVancouverIslandcohosalmonfromlocalCUswerethedominantcontributorstothefisheryForexamplethewestcoastofVancouverIslandCUcontributedapproximately80ofthe39-individualAugustsamplefromtheinshoresouthwesternVancouverIslandregionIntheStraitofGeorgia localCUswereagainthemaincontributorstotherec-reationalfisheryduringthesummerwiththeHoweSoundndashBurrardInletCUcontributingupto40insomesamplestheeastcoastofVancouverIslandCUuptoapproximately30andthelowerFraserRiverCUuptoapproximately40WhilethelowerFraserRiverCUwasanimportantcontributortotherecreationalfisheryintheStraitofGeorgiaandJuandeFucaStraititscontributiontofisheriesoffthewestcoastofVancouverIslandwaslimitedTheNooksackRiverregion innorthernWashingtonStatewasalso identifiedasacon-sistent contributor to recreational fishery samples in the Strait ofGeorgia insomecasescontributingupto20ofthesampleana-lyzed(SupportingInformationTableS2)
StockcompositionswereavailablefromtwotestfisheriesoneatrolltestfisherynearBrooksPeninsulabynorthwesternVancouverIslandandtheotheragillnettestfisheryinupperJohnstoneStraitIn the Brooks Peninsula fishery Vancouver IslandCUs comprised40ndash60ofthesampleswithaminorcontributionofFraserRiverCUs (lt10) The five interior Fraser River CUs of conservationconcern comprised00ndash33of the sampleswhereas themoreabundant lower Fraser River CUs comprised only 00ndash55 ofthesamples Individuals fromFraserRiverCUswerenotdetectedduringJulyinArea127theoffshoreportionofnorthernnorthwest-ernVancouverIslandbuttheyweredetectedinArea126themoresoutherlyoffshoreportionofnorthwesternVancouverIslandIntheJohnstoneStrait(RoundIsland)testfisheryindividualsfromFraserRiverCUswerescarcebutthefiveinteriorFraserRiverCUscom-prised30ndash35of the sampleswhereas the lowerFraserRiverCUscomprisedonly00ndash65ofthesamplesThemajorcontrib-utors to the samplesoriginated from theeast coastofVancouverIsland(24ndash45)andHoweSoundndashBurrardInletCUs(9ndash11)aswellascoastalWashington(8ndash15)(SupportingInformationTableS2)
ThefinalfisherysampledwasademonstrationcommercialtrollfisheryincentralcoastalBC(A6andA7)ContributionsfromlocalCUswerehigherinsamplesfrommoreinshoreareas(A7-7August212southernCUsandregions)withcontributionsfromsouth-ernCUshigherinthemoreseawardareas(A6-9September862southernCUsandregions)LikethewestcoastofVancouverIslandmigrating stocks were more likely to be found in more seawardlocations
emspensp emsp | emsp11BEACHAM Et Al
33emsp|emspEvaluation of 2017 fishery sampling
Onesourceoffisherysampleswasdirectfisherysamplingofindi-viduals(bothadiposefin-clippedandunclipped)wherethefocuswas not associated with potential CWT recovery In this casesamplingofunclippedindividualsfromthenorthern(AreaF)trollfishery indicated that initially (July 10ndash12) northern and centralBCCUscontributed23tothecatchsouthernBCCUs53andsouthern US regions 23 (Supporting Information Table S3)Stock composition changed markedly in samples collected July13ndash18 with Alaska contributing 12 northern and central BCCUs contributing 72 and southern BCCUs contributing 16withnosouthernUScontributionSamplescollectedduringJuly22ndash28revealedadominantcontributionfromnorthernandcen-tralBCCUs(96)andCUsfromthisregioncontributedbyfarthelargestportionof the catch in subsequent sampleswith south-ernBCCUsandsouthernUSregionslargelyabsent(SupportingInformation Table S3) The average contribution to the weeklysamples from July 22 onwardswas 17 southeast Alaska 16HaidaGwaiindashGrahamIslandLowlandsCU15DouglasChannelndashKitimatArmCU 12Hecate StraitMainlandCU 8NorthernCoastalStreamsCUand7LowerNassCUNotunexpectedlysampling of only clipped individuals from the fishery indicatedverysubstantialdifferencesinstockcompositionbetweenclippedandunclippedindividualsTheaveragecontributiontotheweeklysamples of clipped individuals from July 19 onwards was 32coastalWashington11mid-PugetSound10westVancouverIsland CU and 5 for each of northern Puget Sound and Juande Fuca Strait (Supporting Information Table S4) However theobservedcliprateinthefisherywas16sothecatchwasdomi-natedbyunclippedindividualsTheSeptember2ndash8samplefromthe Haida Gwaii recreational fishery near Langara displayed astockcompositionof42HaidaGwaiindashGrahamIslandLowlandsCU15HecateStraitMainlandCU10LowerSkeenaCUand8 Northern Coastal Streams CU similar to that observed inthe troll fisherywith the local CU (HaidaGwaiindashGraham IslandLowlands) inhigherproportionsaswouldbeexpectedinamoreinshorefishery(SupportingInformationTableS3)
Directsamplingof thecentralcoast troll fisheryrevealedthatonaverage theDouglasChannelndashKitimatArmCUwasthe largestcontributortothefishery(24)followedbytheNorthernCoastalStreamsCU(19)BellaCoolandashDeanRivers(18)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(10) (Supporting InformationTableS3)withallCUsproximaltothelocationofthefisherySamplesfromtherecreationalcentralcoastfisheryindicatedthattheBellaCoolandashDeanRiversCUwasthedominant contributor to the fishery (32) followedby theRiversInletCU(19)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(15)andDouglasChannelndashKitimatArmCU(7) (SupportingInformationTableS3)GiventhelocationofthefisherydominantcontributionsfromtheseCUsweretobeexpected
AsnotedpreviouslysampleswerealsoavailablefromvirtuallyallindividualssampledfromcommercialrecreationalandFirstNationsfisheriesthatwouldberoutinelyprocessedforCWTrecoveryinayearbyacentrallaboratoryIndividualheadsfromcommercialfish-eries sent to the central laboratoryhadbeenpreviously screenedelectronically for thepresenceofaCWTVoluntaryheadsamplesfromadiposefin-clippedindividualsfromrecreationalfisheriesweresenttothecentrallaboratorybutmanyoftheseindividualsdidnotcontainaCWTastherewasnopriorscreeningof these individu-als Stock composition of clipped individuals in the northern trollfishery has been previously outlined above Stock composition inthenorthern recreational fishery indicatedawidegeographicdis-tributionofcontributions rangingfromcoastalWashington (16)thewestcoastofVancouverIslandCU(10)theupperSkeenaCU(9)southeastAlaskaregion(8)mid-PugetSound(7)andeastcoast of Vancouver IslandndashStrait of Georgia CU (6) (SupportingInformation Table S4) Stock composition of clipped individuals inthecentralcoastrecreationalfisherywasdominatedbytheHoweSoundndashBurrardInletCU(27)andeastVancouverIslandndashStraitofGeorgiaCU(25)(SupportingInformationTableS4)butgiventhelowobservedclip rate in the fishery (014) therewasvery littlecatchthatcouldbeidentifiedashatcheryorigin
Directsamplingofrecreationalfisheries insouthernBCwhichincluded both clipped and unclipped individuals in the sample
Population Genotyped Assigned assignment
accuracy of population assignment
Robertson 66 66 1000 1000
Quinsam 69 68 986 1000
Puntledge 21 19 905 1000
BigQualicum 84 84 1000 1000
Goldstream 27 19 704 1000
Chilliwack 195 183 938 1000
Inch 91 85 934 1000
Stave 20 19 950 1000
Total 573 543 948 1000
TA B L E 3 emspPercentageofadiposefin-clippedjacks(age2years)sampledin2016andassignedviaparentageanalysistoparentsin2014broodstockpopulationsPopulationofsamplingwasassumedtobethepopulationoforiginofthejacksforevaluationofaccuracyofassignment
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp7BEACHAM Et Al
individualsweredistributedoneachP1chipwith1536individualsprocessedbetweentwochipsThechipswerethenloadedontotheIonTorrentProtonsequencerAfter thesequencingrunwascom-pletedcomparisonswiththereferencegenomeoftherainbowtrout(O mykiss)(Berthelotetal2014)supplementedwiththesequencescontaining theobservedcohosalmonSNPswereconductedwithProton softwareVariant Callerreg and SNP genotypes at the sitesspecified by the hotspot filewithin target regionswere called byVariantCallerThehotspotfilecontained304SNPsiteswithoneSNPscoredateachampliconGenotypesatallavailableSNPsforanindividualwereassembled toprovideamulti-locus individualgen-otype The species identification SNPOkiOts_120255‐113 (StarksClementoampGarza2016)andsex identificationSNPOts_SEXY3‐1 wereomittedfromsubsequentparentageandGSIanalysesleaving302SNPs
26emsp|emspIdentification of individuals
AsnotedpreviouslyPBTandGSIwereusedconcurrently toesti-mate fishery stock composition Initially PBT was used and theanalysiswas conductedwhere the genotypes of individuals to beidentifiedwerematched to the genotypes of prospective parents(COLONY JonesampWang 2010Wang 2016) If all individuals ina hatchery broodstock are sampled and subsequently genotypedthenalloffspringfromthebroodstockaregeneticallymarkedThegenotypes of individuals of unknown origin are statistically com-pared with the genotypes of potential parents and if a match ismadetheoffspringareassignedtotheparentsandthustheori-ginandageoftheindividualsaredeterminedParentageassignmentsoftwarewasutilizedtoassignoffspringtoparentsasCOLONYcanproduceassignmentswhen thegenotypeofoneof theparents ismissingeitherduetoamissingparentalsampleor failure topro-duceaparentalgenotypefromanexistingsampleGiventhatPBTassignments for20potential populationswereevaluated for eachfisherysampleCOLONYwasrunwithallbroodstocksampleddur-ing2014inputasasingleunitforanalysisoffisherysampleswithnodifferentiationamongpopulationsAlthoughtheCOLONYassump-tionofasinglepopulationintheparentpoolwasviolatedanalysisofknown-originsamplesindicatedthatveryhighlevelsofaccuracywereachievedinassignmentswhenpoolingofpotentialparentsincontributing populationswas conducted Two-parent assignmentswere accepted only when both assigned parents originated fromthesamepopulationotherwisetheindividualwaspassedtogeneticstockidentification(GSI)forpotentialassignmentTwo-parentandsingle-parentassignmentswereacceptedonlywhentheprobabilityofcorrectassignmentwasge085fortheparentpairotherwisetheindividualwaspassedtopotentialassignmentbyGSIAdditionallyforsingle-parentassignments tobeaccepted thePBTassignmenttopopulationhad tobepart of theCUassigned to the individualviaGSI Individuals forwhichnoprospectiveparentswere identi-fiedinthebroodstockwerepassedtoGSIforpotentialassignmentPolygamousmatingwasassumedfortheCOLONYanalysisSimplepairwisecomparisonsbetweenoffspringandpotentialparentswere
conductedThebaselineforindividualssampledinthe2016escape-ments (jacks)and2017fisheries includedallbroodstockssampledin2014as these individualsarepredominately threeyearsofage(Sandercock1991)Jacks inthe2017escapementwere identifiedvia body size and subsequent assignment to parents in the 2015hatcherybroodstocksIndividualswithmorethan120missinggeno-typeswereeliminatedfromfurtheranalysesAnestimatedgenotyp-ingerrorrateof1wasusedforCOLONYassignmentsPreviouslyBeachametal(2017)hadreportedthatanaveragegenotypingerrorrate of 107 (1220discrepancies in 114105 comparisons) or analleleerrorrateof053(1220discrepanciesin228210compari-sons)wasobservedoverthe302SNPsscoredTheparentpairout-putfilewasthebasicfileusedinsubsequentanalyses
Thesecondmethodof individual identification isGSI inwhichthegeneticprofilesofwholepopulationspotentiallycontributingtoamixed-stocksampleareusedtoestimatetheoriginofeachindivid-ualinthesample(RUBIASMoranampAnderson2018)ThisanalysiswasrestrictedtothoseindividualsunassignedviaCOLONYForeachsample individuals not assigned by COLONYwere then assignedwithRUBIASThepopulationposteriormeansfilewasthebasicfileusedforsubsequentanalysesandthisfilecontainedtheprobabilityofassignmentoftheindividualtoeachofthe267populationsinthebaselineStockcompositionwasestimatedthroughthecombinationoffilesgeneratedwithbothCOLONYandRUBIAS Individualsas-signedviaCOLONYwereassignedaprobabilityof100oforiginat-ingfromtheidentifiedpopulationwitha000probabilityassignedtoallotherpopulationsinthebaselineThislevelofassignmentac-curacyviaPBTwasobservedpreviouslyforcohosalmonwiththepanelofSNPsemployedinthecurrentstudy(Beachametal2017)ThesedatawerethencombinedwiththeprobabilityofassignmentforthoseindividualsunassignedviaCOLONYtoeachofthepopu-lationsinthebaselineviaRUBIASAfteraninitialburn-inof25000iterations the last1000 iterations from theMonteCarloMarkovchainfromRUBIASwereusedtoestimatetheoriginofindividualsandstockcompositionwiththemeanallocationtoeachpopulationinthebaselinedeterminedStandarddeviationsofestimatedstockcompositionswere alsodetermined from the last1000 iterationsfrom theMonteCarloMarkovchain Stock compositionbyCUorreportinggroupwasdeterminedbysummationofallocationstoallpopulations in the baseline that belonged to the CU or reportinggroupunderconsideration
3emsp |emspRESULTS
31emsp|emspEstimation of stock composition for known‐origin samples
Genotypeswere available from573 jacks sampled in2016acrossthreeCUs and eight populations in southernBC Estimated stockcompositionbyCUand reportinggroup for this combinedsamplewas accuratewith an error utilizingonlyGSI of le04byCU forthreeCUspresentinthesample(Table2)WithbothPBTandGSIutilized theaverageerrordeclinedtole02byCUfor threeCUs
8emsp |emsp emspensp BEACHAM Et Al
TAB
LE 2
emspAccuracyofregional(UnitedStates)andconservationunit(Canada)estimatedstockcomposition()foramixed-stocksampleof573known-origincohosalmonjackssampledin
2016and620coded-wiretaggedindividualssampledin2017fisheriesinBCStandarddeviationoftheestimateisinparentheses
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
SoutheastAlaska
00
00(00)
00(00)
90
68(11)
68(11)
AlsekRiver
00
00(00)
00(00)
00
00(00)
00(00)
LowerStikine
00
00(00)
00(00)
00
04(03)
04(03)
LowerNass
00
00(00)
00(00)
63
64(10)
62(10)
UpperNass
00
00(00)
00(00)
00
04(03)
04(04)
PortlandSound-Observatory
Inlet-PortlandCanal
00
00(00)
00(00)
00
00(00)
00(00)
SkeenaEstuary
00
00(00)
00(00)
00
00(00)
00(00)
LowerSkeena
00
00(00)
00(00)
16
12(05)
12(05)
MiddleSkeena
00
00(00)
00(00)
69
39(08)
39(09)
UpperSkeena
00
00(00)
00(00)
06
40(10)
40(09)
HaidaGwaiindashGrahamIslandLowlands
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-East
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-W
est
00
00(00)
00(00)
00
00(00)
00(00)
NorthernCoastalStreams
00
00(00)
00(00)
00
00(00)
00(01)
HecateStraitMainland
00
00(00)
00(00)
00
07(05)
06(04)
MusselndashKynoch
00
00(00)
00(00)
00
00(00)
00(00)
DouglasChannelndashKitimatArm
00
00(01)
00(01)
00
00(01)
00(01)
BellaCoolandashDeanRivers
00
00(00)
00(00)
00
00(00)
00(00)
RiversInlet
00
00(01)
00(00)
00
00(00)
00(00)
SmithInlet
00
00(00)
00(00)
00
00(00)
00(00)
SouthernCoastalStreamsndashQueen
CharlotteStraitndashJohnstoneStraitndash
SouthernFjords
00
00(01)
00(01)
00
00(02)
00(02)
HomathkondashKlinakliniRivers
00
00(00)
00(00)
00
00(00)
00(00)
GeorgiaStraitMainland
00
00(00)
00(00)
00
00(00)
00(00)
HoweSoundndashBurrardInlet
00
00(00)
00(00)
06
00(01)
00(00)
EastVancouverIslandndashGeorgiaStrait
351
345(20)
348(20)
131
134(14)
140(14)
EastVancouverIslandndashJohnstone
StraitndashSouthernFjords
00
00(00)
00(00)
00
00(01)
00(00)
NahwittiLowland
00
00(01)
00(00)
18
17(06)
17(06) (C
ontin
ues)
emspensp emsp | emsp9BEACHAM Et Al
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
WestVancouverIsland
115
115(14)
115(13)
135
134(14)
135(14)
Clayoquot
00
00(00)
00(00)
00
02(03)
02(02)
JuandeFucandashPachena
00
00(01)
00(01)
00
00(00)
00(00)
LowerFraser
534
539(21)
536(21)
185
187(15)
188(15)
Lillooet
00
00(00)
00(00)
00
03(02)
02(01)
FraserCanyon
00
00(00)
00(00)
00
00(00)
00(00)
InteriorFraser
00
00(00)
00(00)
00
00(00)
00(00)
LowerThompson
00
00(00)
00(00)
08
07(03)
08(04)
NorthThompson
00
00(00)
00(01)
00
00(00)
00(00)
SouthThompson
00
00(00)
00(00)
05
06(03)
05(03)
Boun
dary
Bay
00
00(00)
00(00)
00
00(00)
00(00)
NooksackRiver
00
01(02)
01(02)
26
09(09)
10(08)
SkagitRiver
00
00(01)
00(01)
05
23(08)
22(08)
NorthernPugetSound
00
00(00)
00(00)
39
58(12)
55(13)
Mid-PugetSound
00
00(00)
00(00)
52
62(11)
63(11)
SouthernPugetSound
00
00(00)
00(00)
10
09(10)
06(08)
JuandeFucaStrait
00
00(00)
00(00)
00
02(03)
01(03)
HoodCanal
00
00(00)
00(00)
44
16(05)
16(06)
CoastalWashington
00
00(00)
00(00)
60
59(10)
59(10)
ColumbiaRiver
00
00(00)
00(00)
32
32(07)
32(07)
Oregon
00
00(00)
00(00)
00
00(01)
00(00)
TAB
LE 2
(Continued)
10emsp |emsp emspensp BEACHAM Et Al
presentinthesample(Table2)Theaverageerrorfor45CUsorre-portinggroupsabsentinthesamplewas00bybothGSIandPBT-GSIBypopulation543(948)ofthejackswereassignedviaPBTwith 100 accuracywith respect to population of origin and age(Table3)Estimatedstockcompositionfor20populationswhereitwaspossibletousebothPBTandGSIinestimationofstockcompo-sitionwasaccurateFortheeightpopulationspresentinthesampleofjackstheaverageerrorutilizingGSIonlywas05perpopulationdecliningto02utilizingbothPBTandGSIundertheassumptionthatthecollectionlocationwasanaccuratereflectionofjackoriginTherewasanaverage00errorforthe12populationswithnorep-resentation inthesamplewithutilizationofeitherGSIorPBT-GSI(Table4)ThelargesterrorutilizingbothPBTandGSI(underestima-tionby06ofactualvalue)wasobservedintheGoldstreamRiverpopulationwhichalsodisplayedthe lowestpercentage (704)ofjacksidentifiedviaPBT(Table3)PBTwhencombinedwithGSIpro-ducedmoreaccurateestimatesofCU-specificandpopulation-spe-cificstockcompositionthanavailablewithonlyGSI
Of the 3054 adipose fin-clipped coho salmon samples re-ceivedforpotentialgenotypingfromacentrallaboratorywhere2017fisherysampleswereprocessedforpotentialCWTrecov-erygenotypeswereobtained for2533 (829)of the individ-uals processed There were 750 CWTs recovered from 3054headsor snouts examined and genotypeswereobtained from622 (829) of the individuals that carried a CWT GenotypesofcohosalmonwithoutCWTsbutsentfromtheCWTheadre-covery laboratorywereobtainedfrom1911of2304 individu-als (829) Release information associatedwith theCWTwasavailable for 620 of the 622CWTs associatedwith genotypedindividuals and these 620 individuals constituted a sample ofknown origin There were individuals from 20 regions or CUspresentinthissampleandforthese20regionsorCUstheav-erageerrorofestimationutilizingonlyGSIwas09perregionorCUand09forbothPBTandGSIcombined(Table2)CWTswererecoveredoriginatingfromsevenpopulationswhereitwaspossibletoincorporatebothGSIandPBTinestimationofstockcompositionFor these sevenpopulations theaverageerror inestimatedstockcompositionsperpopulationwas20withGSIand 03 with PBT-GSI (Table 4) There was an average 00errorforthe13populationswithnorepresentationinthesam-ple with utilization of either GSI or PBT-GSI (Table 4) Similartothesampleof jacksPBTwhencombinedwithGSIproducedmore accurate estimates of population-specific stock composi-tionthanavailablewithonlyGSI
32emsp|emspEvaluation of 2016 fishery sampling
Asbroodstocksamplingatselectedhatcheries insouthernBCdidnot commence until 2014 it was not possible to obtain PBT as-signments for any fishery samples collected in 2016 HoweverGSI analysis of individuals derived from recreational fishery sam-ples intheStraitofGeorgiaJuandeFucaStraitandoffthewestcoastofVancouverIslandinsouthernBCindicatedthattherewas
virtually no contribution from regions orCUsnorthofVancouverIsland (Supporting Information Table S2) The only exceptionwasin thenorthwestcoastofVancouver Islandboth inshoreandoff-shorewhereupto12ofthesamplewasidentifiedasoriginatingfromnorthernandcentralBCCUsUS-origincohosalmonwerees-timatedtohavecontributedapproximately70ofsummerrecrea-tionalsamplesderivedfromoffshorefisheriesalongthewestcoastofVancouverIslandaswellasintheentrancetoJuandeFucaStrait(A21A121)InmoreinshoreareasalongthewestcoastofVancouverIslandcohosalmonfromlocalCUswerethedominantcontributorstothefisheryForexamplethewestcoastofVancouverIslandCUcontributedapproximately80ofthe39-individualAugustsamplefromtheinshoresouthwesternVancouverIslandregionIntheStraitofGeorgia localCUswereagainthemaincontributorstotherec-reationalfisheryduringthesummerwiththeHoweSoundndashBurrardInletCUcontributingupto40insomesamplestheeastcoastofVancouverIslandCUuptoapproximately30andthelowerFraserRiverCUuptoapproximately40WhilethelowerFraserRiverCUwasanimportantcontributortotherecreationalfisheryintheStraitofGeorgiaandJuandeFucaStraititscontributiontofisheriesoffthewestcoastofVancouverIslandwaslimitedTheNooksackRiverregion innorthernWashingtonStatewasalso identifiedasacon-sistent contributor to recreational fishery samples in the Strait ofGeorgia insomecasescontributingupto20ofthesampleana-lyzed(SupportingInformationTableS2)
StockcompositionswereavailablefromtwotestfisheriesoneatrolltestfisherynearBrooksPeninsulabynorthwesternVancouverIslandandtheotheragillnettestfisheryinupperJohnstoneStraitIn the Brooks Peninsula fishery Vancouver IslandCUs comprised40ndash60ofthesampleswithaminorcontributionofFraserRiverCUs (lt10) The five interior Fraser River CUs of conservationconcern comprised00ndash33of the sampleswhereas themoreabundant lower Fraser River CUs comprised only 00ndash55 ofthesamples Individuals fromFraserRiverCUswerenotdetectedduringJulyinArea127theoffshoreportionofnorthernnorthwest-ernVancouverIslandbuttheyweredetectedinArea126themoresoutherlyoffshoreportionofnorthwesternVancouverIslandIntheJohnstoneStrait(RoundIsland)testfisheryindividualsfromFraserRiverCUswerescarcebutthefiveinteriorFraserRiverCUscom-prised30ndash35of the sampleswhereas the lowerFraserRiverCUscomprisedonly00ndash65ofthesamplesThemajorcontrib-utors to the samplesoriginated from theeast coastofVancouverIsland(24ndash45)andHoweSoundndashBurrardInletCUs(9ndash11)aswellascoastalWashington(8ndash15)(SupportingInformationTableS2)
ThefinalfisherysampledwasademonstrationcommercialtrollfisheryincentralcoastalBC(A6andA7)ContributionsfromlocalCUswerehigherinsamplesfrommoreinshoreareas(A7-7August212southernCUsandregions)withcontributionsfromsouth-ernCUshigherinthemoreseawardareas(A6-9September862southernCUsandregions)LikethewestcoastofVancouverIslandmigrating stocks were more likely to be found in more seawardlocations
emspensp emsp | emsp11BEACHAM Et Al
33emsp|emspEvaluation of 2017 fishery sampling
Onesourceoffisherysampleswasdirectfisherysamplingofindi-viduals(bothadiposefin-clippedandunclipped)wherethefocuswas not associated with potential CWT recovery In this casesamplingofunclippedindividualsfromthenorthern(AreaF)trollfishery indicated that initially (July 10ndash12) northern and centralBCCUscontributed23tothecatchsouthernBCCUs53andsouthern US regions 23 (Supporting Information Table S3)Stock composition changed markedly in samples collected July13ndash18 with Alaska contributing 12 northern and central BCCUs contributing 72 and southern BCCUs contributing 16withnosouthernUScontributionSamplescollectedduringJuly22ndash28revealedadominantcontributionfromnorthernandcen-tralBCCUs(96)andCUsfromthisregioncontributedbyfarthelargestportionof the catch in subsequent sampleswith south-ernBCCUsandsouthernUSregionslargelyabsent(SupportingInformation Table S3) The average contribution to the weeklysamples from July 22 onwardswas 17 southeast Alaska 16HaidaGwaiindashGrahamIslandLowlandsCU15DouglasChannelndashKitimatArmCU 12Hecate StraitMainlandCU 8NorthernCoastalStreamsCUand7LowerNassCUNotunexpectedlysampling of only clipped individuals from the fishery indicatedverysubstantialdifferencesinstockcompositionbetweenclippedandunclippedindividualsTheaveragecontributiontotheweeklysamples of clipped individuals from July 19 onwards was 32coastalWashington11mid-PugetSound10westVancouverIsland CU and 5 for each of northern Puget Sound and Juande Fuca Strait (Supporting Information Table S4) However theobservedcliprateinthefisherywas16sothecatchwasdomi-natedbyunclippedindividualsTheSeptember2ndash8samplefromthe Haida Gwaii recreational fishery near Langara displayed astockcompositionof42HaidaGwaiindashGrahamIslandLowlandsCU15HecateStraitMainlandCU10LowerSkeenaCUand8 Northern Coastal Streams CU similar to that observed inthe troll fisherywith the local CU (HaidaGwaiindashGraham IslandLowlands) inhigherproportionsaswouldbeexpectedinamoreinshorefishery(SupportingInformationTableS3)
Directsamplingof thecentralcoast troll fisheryrevealedthatonaverage theDouglasChannelndashKitimatArmCUwasthe largestcontributortothefishery(24)followedbytheNorthernCoastalStreamsCU(19)BellaCoolandashDeanRivers(18)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(10) (Supporting InformationTableS3)withallCUsproximaltothelocationofthefisherySamplesfromtherecreationalcentralcoastfisheryindicatedthattheBellaCoolandashDeanRiversCUwasthedominant contributor to the fishery (32) followedby theRiversInletCU(19)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(15)andDouglasChannelndashKitimatArmCU(7) (SupportingInformationTableS3)GiventhelocationofthefisherydominantcontributionsfromtheseCUsweretobeexpected
AsnotedpreviouslysampleswerealsoavailablefromvirtuallyallindividualssampledfromcommercialrecreationalandFirstNationsfisheriesthatwouldberoutinelyprocessedforCWTrecoveryinayearbyacentrallaboratoryIndividualheadsfromcommercialfish-eries sent to the central laboratoryhadbeenpreviously screenedelectronically for thepresenceofaCWTVoluntaryheadsamplesfromadiposefin-clippedindividualsfromrecreationalfisheriesweresenttothecentrallaboratorybutmanyoftheseindividualsdidnotcontainaCWTastherewasnopriorscreeningof these individu-als Stock composition of clipped individuals in the northern trollfishery has been previously outlined above Stock composition inthenorthern recreational fishery indicatedawidegeographicdis-tributionofcontributions rangingfromcoastalWashington (16)thewestcoastofVancouverIslandCU(10)theupperSkeenaCU(9)southeastAlaskaregion(8)mid-PugetSound(7)andeastcoast of Vancouver IslandndashStrait of Georgia CU (6) (SupportingInformation Table S4) Stock composition of clipped individuals inthecentralcoastrecreationalfisherywasdominatedbytheHoweSoundndashBurrardInletCU(27)andeastVancouverIslandndashStraitofGeorgiaCU(25)(SupportingInformationTableS4)butgiventhelowobservedclip rate in the fishery (014) therewasvery littlecatchthatcouldbeidentifiedashatcheryorigin
Directsamplingofrecreationalfisheries insouthernBCwhichincluded both clipped and unclipped individuals in the sample
Population Genotyped Assigned assignment
accuracy of population assignment
Robertson 66 66 1000 1000
Quinsam 69 68 986 1000
Puntledge 21 19 905 1000
BigQualicum 84 84 1000 1000
Goldstream 27 19 704 1000
Chilliwack 195 183 938 1000
Inch 91 85 934 1000
Stave 20 19 950 1000
Total 573 543 948 1000
TA B L E 3 emspPercentageofadiposefin-clippedjacks(age2years)sampledin2016andassignedviaparentageanalysistoparentsin2014broodstockpopulationsPopulationofsamplingwasassumedtobethepopulationoforiginofthejacksforevaluationofaccuracyofassignment
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
8emsp |emsp emspensp BEACHAM Et Al
TAB
LE 2
emspAccuracyofregional(UnitedStates)andconservationunit(Canada)estimatedstockcomposition()foramixed-stocksampleof573known-origincohosalmonjackssampledin
2016and620coded-wiretaggedindividualssampledin2017fisheriesinBCStandarddeviationoftheestimateisinparentheses
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
SoutheastAlaska
00
00(00)
00(00)
90
68(11)
68(11)
AlsekRiver
00
00(00)
00(00)
00
00(00)
00(00)
LowerStikine
00
00(00)
00(00)
00
04(03)
04(03)
LowerNass
00
00(00)
00(00)
63
64(10)
62(10)
UpperNass
00
00(00)
00(00)
00
04(03)
04(04)
PortlandSound-Observatory
Inlet-PortlandCanal
00
00(00)
00(00)
00
00(00)
00(00)
SkeenaEstuary
00
00(00)
00(00)
00
00(00)
00(00)
LowerSkeena
00
00(00)
00(00)
16
12(05)
12(05)
MiddleSkeena
00
00(00)
00(00)
69
39(08)
39(09)
UpperSkeena
00
00(00)
00(00)
06
40(10)
40(09)
HaidaGwaiindashGrahamIslandLowlands
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-East
00
00(00)
00(00)
00
00(00)
00(00)
HaidaGwaii-W
est
00
00(00)
00(00)
00
00(00)
00(00)
NorthernCoastalStreams
00
00(00)
00(00)
00
00(00)
00(01)
HecateStraitMainland
00
00(00)
00(00)
00
07(05)
06(04)
MusselndashKynoch
00
00(00)
00(00)
00
00(00)
00(00)
DouglasChannelndashKitimatArm
00
00(01)
00(01)
00
00(01)
00(01)
BellaCoolandashDeanRivers
00
00(00)
00(00)
00
00(00)
00(00)
RiversInlet
00
00(01)
00(00)
00
00(00)
00(00)
SmithInlet
00
00(00)
00(00)
00
00(00)
00(00)
SouthernCoastalStreamsndashQueen
CharlotteStraitndashJohnstoneStraitndash
SouthernFjords
00
00(01)
00(01)
00
00(02)
00(02)
HomathkondashKlinakliniRivers
00
00(00)
00(00)
00
00(00)
00(00)
GeorgiaStraitMainland
00
00(00)
00(00)
00
00(00)
00(00)
HoweSoundndashBurrardInlet
00
00(00)
00(00)
06
00(01)
00(00)
EastVancouverIslandndashGeorgiaStrait
351
345(20)
348(20)
131
134(14)
140(14)
EastVancouverIslandndashJohnstone
StraitndashSouthernFjords
00
00(00)
00(00)
00
00(01)
00(00)
NahwittiLowland
00
00(01)
00(00)
18
17(06)
17(06) (C
ontin
ues)
emspensp emsp | emsp9BEACHAM Et Al
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
WestVancouverIsland
115
115(14)
115(13)
135
134(14)
135(14)
Clayoquot
00
00(00)
00(00)
00
02(03)
02(02)
JuandeFucandashPachena
00
00(01)
00(01)
00
00(00)
00(00)
LowerFraser
534
539(21)
536(21)
185
187(15)
188(15)
Lillooet
00
00(00)
00(00)
00
03(02)
02(01)
FraserCanyon
00
00(00)
00(00)
00
00(00)
00(00)
InteriorFraser
00
00(00)
00(00)
00
00(00)
00(00)
LowerThompson
00
00(00)
00(00)
08
07(03)
08(04)
NorthThompson
00
00(00)
00(01)
00
00(00)
00(00)
SouthThompson
00
00(00)
00(00)
05
06(03)
05(03)
Boun
dary
Bay
00
00(00)
00(00)
00
00(00)
00(00)
NooksackRiver
00
01(02)
01(02)
26
09(09)
10(08)
SkagitRiver
00
00(01)
00(01)
05
23(08)
22(08)
NorthernPugetSound
00
00(00)
00(00)
39
58(12)
55(13)
Mid-PugetSound
00
00(00)
00(00)
52
62(11)
63(11)
SouthernPugetSound
00
00(00)
00(00)
10
09(10)
06(08)
JuandeFucaStrait
00
00(00)
00(00)
00
02(03)
01(03)
HoodCanal
00
00(00)
00(00)
44
16(05)
16(06)
CoastalWashington
00
00(00)
00(00)
60
59(10)
59(10)
ColumbiaRiver
00
00(00)
00(00)
32
32(07)
32(07)
Oregon
00
00(00)
00(00)
00
00(01)
00(00)
TAB
LE 2
(Continued)
10emsp |emsp emspensp BEACHAM Et Al
presentinthesample(Table2)Theaverageerrorfor45CUsorre-portinggroupsabsentinthesamplewas00bybothGSIandPBT-GSIBypopulation543(948)ofthejackswereassignedviaPBTwith 100 accuracywith respect to population of origin and age(Table3)Estimatedstockcompositionfor20populationswhereitwaspossibletousebothPBTandGSIinestimationofstockcompo-sitionwasaccurateFortheeightpopulationspresentinthesampleofjackstheaverageerrorutilizingGSIonlywas05perpopulationdecliningto02utilizingbothPBTandGSIundertheassumptionthatthecollectionlocationwasanaccuratereflectionofjackoriginTherewasanaverage00errorforthe12populationswithnorep-resentation inthesamplewithutilizationofeitherGSIorPBT-GSI(Table4)ThelargesterrorutilizingbothPBTandGSI(underestima-tionby06ofactualvalue)wasobservedintheGoldstreamRiverpopulationwhichalsodisplayedthe lowestpercentage (704)ofjacksidentifiedviaPBT(Table3)PBTwhencombinedwithGSIpro-ducedmoreaccurateestimatesofCU-specificandpopulation-spe-cificstockcompositionthanavailablewithonlyGSI
Of the 3054 adipose fin-clipped coho salmon samples re-ceivedforpotentialgenotypingfromacentrallaboratorywhere2017fisherysampleswereprocessedforpotentialCWTrecov-erygenotypeswereobtained for2533 (829)of the individ-uals processed There were 750 CWTs recovered from 3054headsor snouts examined and genotypeswereobtained from622 (829) of the individuals that carried a CWT GenotypesofcohosalmonwithoutCWTsbutsentfromtheCWTheadre-covery laboratorywereobtainedfrom1911of2304 individu-als (829) Release information associatedwith theCWTwasavailable for 620 of the 622CWTs associatedwith genotypedindividuals and these 620 individuals constituted a sample ofknown origin There were individuals from 20 regions or CUspresentinthissampleandforthese20regionsorCUstheav-erageerrorofestimationutilizingonlyGSIwas09perregionorCUand09forbothPBTandGSIcombined(Table2)CWTswererecoveredoriginatingfromsevenpopulationswhereitwaspossibletoincorporatebothGSIandPBTinestimationofstockcompositionFor these sevenpopulations theaverageerror inestimatedstockcompositionsperpopulationwas20withGSIand 03 with PBT-GSI (Table 4) There was an average 00errorforthe13populationswithnorepresentationinthesam-ple with utilization of either GSI or PBT-GSI (Table 4) Similartothesampleof jacksPBTwhencombinedwithGSIproducedmore accurate estimates of population-specific stock composi-tionthanavailablewithonlyGSI
32emsp|emspEvaluation of 2016 fishery sampling
Asbroodstocksamplingatselectedhatcheries insouthernBCdidnot commence until 2014 it was not possible to obtain PBT as-signments for any fishery samples collected in 2016 HoweverGSI analysis of individuals derived from recreational fishery sam-ples intheStraitofGeorgiaJuandeFucaStraitandoffthewestcoastofVancouverIslandinsouthernBCindicatedthattherewas
virtually no contribution from regions orCUsnorthofVancouverIsland (Supporting Information Table S2) The only exceptionwasin thenorthwestcoastofVancouver Islandboth inshoreandoff-shorewhereupto12ofthesamplewasidentifiedasoriginatingfromnorthernandcentralBCCUsUS-origincohosalmonwerees-timatedtohavecontributedapproximately70ofsummerrecrea-tionalsamplesderivedfromoffshorefisheriesalongthewestcoastofVancouverIslandaswellasintheentrancetoJuandeFucaStrait(A21A121)InmoreinshoreareasalongthewestcoastofVancouverIslandcohosalmonfromlocalCUswerethedominantcontributorstothefisheryForexamplethewestcoastofVancouverIslandCUcontributedapproximately80ofthe39-individualAugustsamplefromtheinshoresouthwesternVancouverIslandregionIntheStraitofGeorgia localCUswereagainthemaincontributorstotherec-reationalfisheryduringthesummerwiththeHoweSoundndashBurrardInletCUcontributingupto40insomesamplestheeastcoastofVancouverIslandCUuptoapproximately30andthelowerFraserRiverCUuptoapproximately40WhilethelowerFraserRiverCUwasanimportantcontributortotherecreationalfisheryintheStraitofGeorgiaandJuandeFucaStraititscontributiontofisheriesoffthewestcoastofVancouverIslandwaslimitedTheNooksackRiverregion innorthernWashingtonStatewasalso identifiedasacon-sistent contributor to recreational fishery samples in the Strait ofGeorgia insomecasescontributingupto20ofthesampleana-lyzed(SupportingInformationTableS2)
StockcompositionswereavailablefromtwotestfisheriesoneatrolltestfisherynearBrooksPeninsulabynorthwesternVancouverIslandandtheotheragillnettestfisheryinupperJohnstoneStraitIn the Brooks Peninsula fishery Vancouver IslandCUs comprised40ndash60ofthesampleswithaminorcontributionofFraserRiverCUs (lt10) The five interior Fraser River CUs of conservationconcern comprised00ndash33of the sampleswhereas themoreabundant lower Fraser River CUs comprised only 00ndash55 ofthesamples Individuals fromFraserRiverCUswerenotdetectedduringJulyinArea127theoffshoreportionofnorthernnorthwest-ernVancouverIslandbuttheyweredetectedinArea126themoresoutherlyoffshoreportionofnorthwesternVancouverIslandIntheJohnstoneStrait(RoundIsland)testfisheryindividualsfromFraserRiverCUswerescarcebutthefiveinteriorFraserRiverCUscom-prised30ndash35of the sampleswhereas the lowerFraserRiverCUscomprisedonly00ndash65ofthesamplesThemajorcontrib-utors to the samplesoriginated from theeast coastofVancouverIsland(24ndash45)andHoweSoundndashBurrardInletCUs(9ndash11)aswellascoastalWashington(8ndash15)(SupportingInformationTableS2)
ThefinalfisherysampledwasademonstrationcommercialtrollfisheryincentralcoastalBC(A6andA7)ContributionsfromlocalCUswerehigherinsamplesfrommoreinshoreareas(A7-7August212southernCUsandregions)withcontributionsfromsouth-ernCUshigherinthemoreseawardareas(A6-9September862southernCUsandregions)LikethewestcoastofVancouverIslandmigrating stocks were more likely to be found in more seawardlocations
emspensp emsp | emsp11BEACHAM Et Al
33emsp|emspEvaluation of 2017 fishery sampling
Onesourceoffisherysampleswasdirectfisherysamplingofindi-viduals(bothadiposefin-clippedandunclipped)wherethefocuswas not associated with potential CWT recovery In this casesamplingofunclippedindividualsfromthenorthern(AreaF)trollfishery indicated that initially (July 10ndash12) northern and centralBCCUscontributed23tothecatchsouthernBCCUs53andsouthern US regions 23 (Supporting Information Table S3)Stock composition changed markedly in samples collected July13ndash18 with Alaska contributing 12 northern and central BCCUs contributing 72 and southern BCCUs contributing 16withnosouthernUScontributionSamplescollectedduringJuly22ndash28revealedadominantcontributionfromnorthernandcen-tralBCCUs(96)andCUsfromthisregioncontributedbyfarthelargestportionof the catch in subsequent sampleswith south-ernBCCUsandsouthernUSregionslargelyabsent(SupportingInformation Table S3) The average contribution to the weeklysamples from July 22 onwardswas 17 southeast Alaska 16HaidaGwaiindashGrahamIslandLowlandsCU15DouglasChannelndashKitimatArmCU 12Hecate StraitMainlandCU 8NorthernCoastalStreamsCUand7LowerNassCUNotunexpectedlysampling of only clipped individuals from the fishery indicatedverysubstantialdifferencesinstockcompositionbetweenclippedandunclippedindividualsTheaveragecontributiontotheweeklysamples of clipped individuals from July 19 onwards was 32coastalWashington11mid-PugetSound10westVancouverIsland CU and 5 for each of northern Puget Sound and Juande Fuca Strait (Supporting Information Table S4) However theobservedcliprateinthefisherywas16sothecatchwasdomi-natedbyunclippedindividualsTheSeptember2ndash8samplefromthe Haida Gwaii recreational fishery near Langara displayed astockcompositionof42HaidaGwaiindashGrahamIslandLowlandsCU15HecateStraitMainlandCU10LowerSkeenaCUand8 Northern Coastal Streams CU similar to that observed inthe troll fisherywith the local CU (HaidaGwaiindashGraham IslandLowlands) inhigherproportionsaswouldbeexpectedinamoreinshorefishery(SupportingInformationTableS3)
Directsamplingof thecentralcoast troll fisheryrevealedthatonaverage theDouglasChannelndashKitimatArmCUwasthe largestcontributortothefishery(24)followedbytheNorthernCoastalStreamsCU(19)BellaCoolandashDeanRivers(18)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(10) (Supporting InformationTableS3)withallCUsproximaltothelocationofthefisherySamplesfromtherecreationalcentralcoastfisheryindicatedthattheBellaCoolandashDeanRiversCUwasthedominant contributor to the fishery (32) followedby theRiversInletCU(19)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(15)andDouglasChannelndashKitimatArmCU(7) (SupportingInformationTableS3)GiventhelocationofthefisherydominantcontributionsfromtheseCUsweretobeexpected
AsnotedpreviouslysampleswerealsoavailablefromvirtuallyallindividualssampledfromcommercialrecreationalandFirstNationsfisheriesthatwouldberoutinelyprocessedforCWTrecoveryinayearbyacentrallaboratoryIndividualheadsfromcommercialfish-eries sent to the central laboratoryhadbeenpreviously screenedelectronically for thepresenceofaCWTVoluntaryheadsamplesfromadiposefin-clippedindividualsfromrecreationalfisheriesweresenttothecentrallaboratorybutmanyoftheseindividualsdidnotcontainaCWTastherewasnopriorscreeningof these individu-als Stock composition of clipped individuals in the northern trollfishery has been previously outlined above Stock composition inthenorthern recreational fishery indicatedawidegeographicdis-tributionofcontributions rangingfromcoastalWashington (16)thewestcoastofVancouverIslandCU(10)theupperSkeenaCU(9)southeastAlaskaregion(8)mid-PugetSound(7)andeastcoast of Vancouver IslandndashStrait of Georgia CU (6) (SupportingInformation Table S4) Stock composition of clipped individuals inthecentralcoastrecreationalfisherywasdominatedbytheHoweSoundndashBurrardInletCU(27)andeastVancouverIslandndashStraitofGeorgiaCU(25)(SupportingInformationTableS4)butgiventhelowobservedclip rate in the fishery (014) therewasvery littlecatchthatcouldbeidentifiedashatcheryorigin
Directsamplingofrecreationalfisheries insouthernBCwhichincluded both clipped and unclipped individuals in the sample
Population Genotyped Assigned assignment
accuracy of population assignment
Robertson 66 66 1000 1000
Quinsam 69 68 986 1000
Puntledge 21 19 905 1000
BigQualicum 84 84 1000 1000
Goldstream 27 19 704 1000
Chilliwack 195 183 938 1000
Inch 91 85 934 1000
Stave 20 19 950 1000
Total 573 543 948 1000
TA B L E 3 emspPercentageofadiposefin-clippedjacks(age2years)sampledin2016andassignedviaparentageanalysistoparentsin2014broodstockpopulationsPopulationofsamplingwasassumedtobethepopulationoforiginofthejacksforevaluationofaccuracyofassignment
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp9BEACHAM Et Al
Regi
onc
onse
rvat
ion
unit
2016
jack
s20
17 C
WT
True
GSI
PBT‐
GSI
True
GSI
PBT‐
GSI
WestVancouverIsland
115
115(14)
115(13)
135
134(14)
135(14)
Clayoquot
00
00(00)
00(00)
00
02(03)
02(02)
JuandeFucandashPachena
00
00(01)
00(01)
00
00(00)
00(00)
LowerFraser
534
539(21)
536(21)
185
187(15)
188(15)
Lillooet
00
00(00)
00(00)
00
03(02)
02(01)
FraserCanyon
00
00(00)
00(00)
00
00(00)
00(00)
InteriorFraser
00
00(00)
00(00)
00
00(00)
00(00)
LowerThompson
00
00(00)
00(00)
08
07(03)
08(04)
NorthThompson
00
00(00)
00(01)
00
00(00)
00(00)
SouthThompson
00
00(00)
00(00)
05
06(03)
05(03)
Boun
dary
Bay
00
00(00)
00(00)
00
00(00)
00(00)
NooksackRiver
00
01(02)
01(02)
26
09(09)
10(08)
SkagitRiver
00
00(01)
00(01)
05
23(08)
22(08)
NorthernPugetSound
00
00(00)
00(00)
39
58(12)
55(13)
Mid-PugetSound
00
00(00)
00(00)
52
62(11)
63(11)
SouthernPugetSound
00
00(00)
00(00)
10
09(10)
06(08)
JuandeFucaStrait
00
00(00)
00(00)
00
02(03)
01(03)
HoodCanal
00
00(00)
00(00)
44
16(05)
16(06)
CoastalWashington
00
00(00)
00(00)
60
59(10)
59(10)
ColumbiaRiver
00
00(00)
00(00)
32
32(07)
32(07)
Oregon
00
00(00)
00(00)
00
00(01)
00(00)
TAB
LE 2
(Continued)
10emsp |emsp emspensp BEACHAM Et Al
presentinthesample(Table2)Theaverageerrorfor45CUsorre-portinggroupsabsentinthesamplewas00bybothGSIandPBT-GSIBypopulation543(948)ofthejackswereassignedviaPBTwith 100 accuracywith respect to population of origin and age(Table3)Estimatedstockcompositionfor20populationswhereitwaspossibletousebothPBTandGSIinestimationofstockcompo-sitionwasaccurateFortheeightpopulationspresentinthesampleofjackstheaverageerrorutilizingGSIonlywas05perpopulationdecliningto02utilizingbothPBTandGSIundertheassumptionthatthecollectionlocationwasanaccuratereflectionofjackoriginTherewasanaverage00errorforthe12populationswithnorep-resentation inthesamplewithutilizationofeitherGSIorPBT-GSI(Table4)ThelargesterrorutilizingbothPBTandGSI(underestima-tionby06ofactualvalue)wasobservedintheGoldstreamRiverpopulationwhichalsodisplayedthe lowestpercentage (704)ofjacksidentifiedviaPBT(Table3)PBTwhencombinedwithGSIpro-ducedmoreaccurateestimatesofCU-specificandpopulation-spe-cificstockcompositionthanavailablewithonlyGSI
Of the 3054 adipose fin-clipped coho salmon samples re-ceivedforpotentialgenotypingfromacentrallaboratorywhere2017fisherysampleswereprocessedforpotentialCWTrecov-erygenotypeswereobtained for2533 (829)of the individ-uals processed There were 750 CWTs recovered from 3054headsor snouts examined and genotypeswereobtained from622 (829) of the individuals that carried a CWT GenotypesofcohosalmonwithoutCWTsbutsentfromtheCWTheadre-covery laboratorywereobtainedfrom1911of2304 individu-als (829) Release information associatedwith theCWTwasavailable for 620 of the 622CWTs associatedwith genotypedindividuals and these 620 individuals constituted a sample ofknown origin There were individuals from 20 regions or CUspresentinthissampleandforthese20regionsorCUstheav-erageerrorofestimationutilizingonlyGSIwas09perregionorCUand09forbothPBTandGSIcombined(Table2)CWTswererecoveredoriginatingfromsevenpopulationswhereitwaspossibletoincorporatebothGSIandPBTinestimationofstockcompositionFor these sevenpopulations theaverageerror inestimatedstockcompositionsperpopulationwas20withGSIand 03 with PBT-GSI (Table 4) There was an average 00errorforthe13populationswithnorepresentationinthesam-ple with utilization of either GSI or PBT-GSI (Table 4) Similartothesampleof jacksPBTwhencombinedwithGSIproducedmore accurate estimates of population-specific stock composi-tionthanavailablewithonlyGSI
32emsp|emspEvaluation of 2016 fishery sampling
Asbroodstocksamplingatselectedhatcheries insouthernBCdidnot commence until 2014 it was not possible to obtain PBT as-signments for any fishery samples collected in 2016 HoweverGSI analysis of individuals derived from recreational fishery sam-ples intheStraitofGeorgiaJuandeFucaStraitandoffthewestcoastofVancouverIslandinsouthernBCindicatedthattherewas
virtually no contribution from regions orCUsnorthofVancouverIsland (Supporting Information Table S2) The only exceptionwasin thenorthwestcoastofVancouver Islandboth inshoreandoff-shorewhereupto12ofthesamplewasidentifiedasoriginatingfromnorthernandcentralBCCUsUS-origincohosalmonwerees-timatedtohavecontributedapproximately70ofsummerrecrea-tionalsamplesderivedfromoffshorefisheriesalongthewestcoastofVancouverIslandaswellasintheentrancetoJuandeFucaStrait(A21A121)InmoreinshoreareasalongthewestcoastofVancouverIslandcohosalmonfromlocalCUswerethedominantcontributorstothefisheryForexamplethewestcoastofVancouverIslandCUcontributedapproximately80ofthe39-individualAugustsamplefromtheinshoresouthwesternVancouverIslandregionIntheStraitofGeorgia localCUswereagainthemaincontributorstotherec-reationalfisheryduringthesummerwiththeHoweSoundndashBurrardInletCUcontributingupto40insomesamplestheeastcoastofVancouverIslandCUuptoapproximately30andthelowerFraserRiverCUuptoapproximately40WhilethelowerFraserRiverCUwasanimportantcontributortotherecreationalfisheryintheStraitofGeorgiaandJuandeFucaStraititscontributiontofisheriesoffthewestcoastofVancouverIslandwaslimitedTheNooksackRiverregion innorthernWashingtonStatewasalso identifiedasacon-sistent contributor to recreational fishery samples in the Strait ofGeorgia insomecasescontributingupto20ofthesampleana-lyzed(SupportingInformationTableS2)
StockcompositionswereavailablefromtwotestfisheriesoneatrolltestfisherynearBrooksPeninsulabynorthwesternVancouverIslandandtheotheragillnettestfisheryinupperJohnstoneStraitIn the Brooks Peninsula fishery Vancouver IslandCUs comprised40ndash60ofthesampleswithaminorcontributionofFraserRiverCUs (lt10) The five interior Fraser River CUs of conservationconcern comprised00ndash33of the sampleswhereas themoreabundant lower Fraser River CUs comprised only 00ndash55 ofthesamples Individuals fromFraserRiverCUswerenotdetectedduringJulyinArea127theoffshoreportionofnorthernnorthwest-ernVancouverIslandbuttheyweredetectedinArea126themoresoutherlyoffshoreportionofnorthwesternVancouverIslandIntheJohnstoneStrait(RoundIsland)testfisheryindividualsfromFraserRiverCUswerescarcebutthefiveinteriorFraserRiverCUscom-prised30ndash35of the sampleswhereas the lowerFraserRiverCUscomprisedonly00ndash65ofthesamplesThemajorcontrib-utors to the samplesoriginated from theeast coastofVancouverIsland(24ndash45)andHoweSoundndashBurrardInletCUs(9ndash11)aswellascoastalWashington(8ndash15)(SupportingInformationTableS2)
ThefinalfisherysampledwasademonstrationcommercialtrollfisheryincentralcoastalBC(A6andA7)ContributionsfromlocalCUswerehigherinsamplesfrommoreinshoreareas(A7-7August212southernCUsandregions)withcontributionsfromsouth-ernCUshigherinthemoreseawardareas(A6-9September862southernCUsandregions)LikethewestcoastofVancouverIslandmigrating stocks were more likely to be found in more seawardlocations
emspensp emsp | emsp11BEACHAM Et Al
33emsp|emspEvaluation of 2017 fishery sampling
Onesourceoffisherysampleswasdirectfisherysamplingofindi-viduals(bothadiposefin-clippedandunclipped)wherethefocuswas not associated with potential CWT recovery In this casesamplingofunclippedindividualsfromthenorthern(AreaF)trollfishery indicated that initially (July 10ndash12) northern and centralBCCUscontributed23tothecatchsouthernBCCUs53andsouthern US regions 23 (Supporting Information Table S3)Stock composition changed markedly in samples collected July13ndash18 with Alaska contributing 12 northern and central BCCUs contributing 72 and southern BCCUs contributing 16withnosouthernUScontributionSamplescollectedduringJuly22ndash28revealedadominantcontributionfromnorthernandcen-tralBCCUs(96)andCUsfromthisregioncontributedbyfarthelargestportionof the catch in subsequent sampleswith south-ernBCCUsandsouthernUSregionslargelyabsent(SupportingInformation Table S3) The average contribution to the weeklysamples from July 22 onwardswas 17 southeast Alaska 16HaidaGwaiindashGrahamIslandLowlandsCU15DouglasChannelndashKitimatArmCU 12Hecate StraitMainlandCU 8NorthernCoastalStreamsCUand7LowerNassCUNotunexpectedlysampling of only clipped individuals from the fishery indicatedverysubstantialdifferencesinstockcompositionbetweenclippedandunclippedindividualsTheaveragecontributiontotheweeklysamples of clipped individuals from July 19 onwards was 32coastalWashington11mid-PugetSound10westVancouverIsland CU and 5 for each of northern Puget Sound and Juande Fuca Strait (Supporting Information Table S4) However theobservedcliprateinthefisherywas16sothecatchwasdomi-natedbyunclippedindividualsTheSeptember2ndash8samplefromthe Haida Gwaii recreational fishery near Langara displayed astockcompositionof42HaidaGwaiindashGrahamIslandLowlandsCU15HecateStraitMainlandCU10LowerSkeenaCUand8 Northern Coastal Streams CU similar to that observed inthe troll fisherywith the local CU (HaidaGwaiindashGraham IslandLowlands) inhigherproportionsaswouldbeexpectedinamoreinshorefishery(SupportingInformationTableS3)
Directsamplingof thecentralcoast troll fisheryrevealedthatonaverage theDouglasChannelndashKitimatArmCUwasthe largestcontributortothefishery(24)followedbytheNorthernCoastalStreamsCU(19)BellaCoolandashDeanRivers(18)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(10) (Supporting InformationTableS3)withallCUsproximaltothelocationofthefisherySamplesfromtherecreationalcentralcoastfisheryindicatedthattheBellaCoolandashDeanRiversCUwasthedominant contributor to the fishery (32) followedby theRiversInletCU(19)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(15)andDouglasChannelndashKitimatArmCU(7) (SupportingInformationTableS3)GiventhelocationofthefisherydominantcontributionsfromtheseCUsweretobeexpected
AsnotedpreviouslysampleswerealsoavailablefromvirtuallyallindividualssampledfromcommercialrecreationalandFirstNationsfisheriesthatwouldberoutinelyprocessedforCWTrecoveryinayearbyacentrallaboratoryIndividualheadsfromcommercialfish-eries sent to the central laboratoryhadbeenpreviously screenedelectronically for thepresenceofaCWTVoluntaryheadsamplesfromadiposefin-clippedindividualsfromrecreationalfisheriesweresenttothecentrallaboratorybutmanyoftheseindividualsdidnotcontainaCWTastherewasnopriorscreeningof these individu-als Stock composition of clipped individuals in the northern trollfishery has been previously outlined above Stock composition inthenorthern recreational fishery indicatedawidegeographicdis-tributionofcontributions rangingfromcoastalWashington (16)thewestcoastofVancouverIslandCU(10)theupperSkeenaCU(9)southeastAlaskaregion(8)mid-PugetSound(7)andeastcoast of Vancouver IslandndashStrait of Georgia CU (6) (SupportingInformation Table S4) Stock composition of clipped individuals inthecentralcoastrecreationalfisherywasdominatedbytheHoweSoundndashBurrardInletCU(27)andeastVancouverIslandndashStraitofGeorgiaCU(25)(SupportingInformationTableS4)butgiventhelowobservedclip rate in the fishery (014) therewasvery littlecatchthatcouldbeidentifiedashatcheryorigin
Directsamplingofrecreationalfisheries insouthernBCwhichincluded both clipped and unclipped individuals in the sample
Population Genotyped Assigned assignment
accuracy of population assignment
Robertson 66 66 1000 1000
Quinsam 69 68 986 1000
Puntledge 21 19 905 1000
BigQualicum 84 84 1000 1000
Goldstream 27 19 704 1000
Chilliwack 195 183 938 1000
Inch 91 85 934 1000
Stave 20 19 950 1000
Total 573 543 948 1000
TA B L E 3 emspPercentageofadiposefin-clippedjacks(age2years)sampledin2016andassignedviaparentageanalysistoparentsin2014broodstockpopulationsPopulationofsamplingwasassumedtobethepopulationoforiginofthejacksforevaluationofaccuracyofassignment
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
10emsp |emsp emspensp BEACHAM Et Al
presentinthesample(Table2)Theaverageerrorfor45CUsorre-portinggroupsabsentinthesamplewas00bybothGSIandPBT-GSIBypopulation543(948)ofthejackswereassignedviaPBTwith 100 accuracywith respect to population of origin and age(Table3)Estimatedstockcompositionfor20populationswhereitwaspossibletousebothPBTandGSIinestimationofstockcompo-sitionwasaccurateFortheeightpopulationspresentinthesampleofjackstheaverageerrorutilizingGSIonlywas05perpopulationdecliningto02utilizingbothPBTandGSIundertheassumptionthatthecollectionlocationwasanaccuratereflectionofjackoriginTherewasanaverage00errorforthe12populationswithnorep-resentation inthesamplewithutilizationofeitherGSIorPBT-GSI(Table4)ThelargesterrorutilizingbothPBTandGSI(underestima-tionby06ofactualvalue)wasobservedintheGoldstreamRiverpopulationwhichalsodisplayedthe lowestpercentage (704)ofjacksidentifiedviaPBT(Table3)PBTwhencombinedwithGSIpro-ducedmoreaccurateestimatesofCU-specificandpopulation-spe-cificstockcompositionthanavailablewithonlyGSI
Of the 3054 adipose fin-clipped coho salmon samples re-ceivedforpotentialgenotypingfromacentrallaboratorywhere2017fisherysampleswereprocessedforpotentialCWTrecov-erygenotypeswereobtained for2533 (829)of the individ-uals processed There were 750 CWTs recovered from 3054headsor snouts examined and genotypeswereobtained from622 (829) of the individuals that carried a CWT GenotypesofcohosalmonwithoutCWTsbutsentfromtheCWTheadre-covery laboratorywereobtainedfrom1911of2304 individu-als (829) Release information associatedwith theCWTwasavailable for 620 of the 622CWTs associatedwith genotypedindividuals and these 620 individuals constituted a sample ofknown origin There were individuals from 20 regions or CUspresentinthissampleandforthese20regionsorCUstheav-erageerrorofestimationutilizingonlyGSIwas09perregionorCUand09forbothPBTandGSIcombined(Table2)CWTswererecoveredoriginatingfromsevenpopulationswhereitwaspossibletoincorporatebothGSIandPBTinestimationofstockcompositionFor these sevenpopulations theaverageerror inestimatedstockcompositionsperpopulationwas20withGSIand 03 with PBT-GSI (Table 4) There was an average 00errorforthe13populationswithnorepresentationinthesam-ple with utilization of either GSI or PBT-GSI (Table 4) Similartothesampleof jacksPBTwhencombinedwithGSIproducedmore accurate estimates of population-specific stock composi-tionthanavailablewithonlyGSI
32emsp|emspEvaluation of 2016 fishery sampling
Asbroodstocksamplingatselectedhatcheries insouthernBCdidnot commence until 2014 it was not possible to obtain PBT as-signments for any fishery samples collected in 2016 HoweverGSI analysis of individuals derived from recreational fishery sam-ples intheStraitofGeorgiaJuandeFucaStraitandoffthewestcoastofVancouverIslandinsouthernBCindicatedthattherewas
virtually no contribution from regions orCUsnorthofVancouverIsland (Supporting Information Table S2) The only exceptionwasin thenorthwestcoastofVancouver Islandboth inshoreandoff-shorewhereupto12ofthesamplewasidentifiedasoriginatingfromnorthernandcentralBCCUsUS-origincohosalmonwerees-timatedtohavecontributedapproximately70ofsummerrecrea-tionalsamplesderivedfromoffshorefisheriesalongthewestcoastofVancouverIslandaswellasintheentrancetoJuandeFucaStrait(A21A121)InmoreinshoreareasalongthewestcoastofVancouverIslandcohosalmonfromlocalCUswerethedominantcontributorstothefisheryForexamplethewestcoastofVancouverIslandCUcontributedapproximately80ofthe39-individualAugustsamplefromtheinshoresouthwesternVancouverIslandregionIntheStraitofGeorgia localCUswereagainthemaincontributorstotherec-reationalfisheryduringthesummerwiththeHoweSoundndashBurrardInletCUcontributingupto40insomesamplestheeastcoastofVancouverIslandCUuptoapproximately30andthelowerFraserRiverCUuptoapproximately40WhilethelowerFraserRiverCUwasanimportantcontributortotherecreationalfisheryintheStraitofGeorgiaandJuandeFucaStraititscontributiontofisheriesoffthewestcoastofVancouverIslandwaslimitedTheNooksackRiverregion innorthernWashingtonStatewasalso identifiedasacon-sistent contributor to recreational fishery samples in the Strait ofGeorgia insomecasescontributingupto20ofthesampleana-lyzed(SupportingInformationTableS2)
StockcompositionswereavailablefromtwotestfisheriesoneatrolltestfisherynearBrooksPeninsulabynorthwesternVancouverIslandandtheotheragillnettestfisheryinupperJohnstoneStraitIn the Brooks Peninsula fishery Vancouver IslandCUs comprised40ndash60ofthesampleswithaminorcontributionofFraserRiverCUs (lt10) The five interior Fraser River CUs of conservationconcern comprised00ndash33of the sampleswhereas themoreabundant lower Fraser River CUs comprised only 00ndash55 ofthesamples Individuals fromFraserRiverCUswerenotdetectedduringJulyinArea127theoffshoreportionofnorthernnorthwest-ernVancouverIslandbuttheyweredetectedinArea126themoresoutherlyoffshoreportionofnorthwesternVancouverIslandIntheJohnstoneStrait(RoundIsland)testfisheryindividualsfromFraserRiverCUswerescarcebutthefiveinteriorFraserRiverCUscom-prised30ndash35of the sampleswhereas the lowerFraserRiverCUscomprisedonly00ndash65ofthesamplesThemajorcontrib-utors to the samplesoriginated from theeast coastofVancouverIsland(24ndash45)andHoweSoundndashBurrardInletCUs(9ndash11)aswellascoastalWashington(8ndash15)(SupportingInformationTableS2)
ThefinalfisherysampledwasademonstrationcommercialtrollfisheryincentralcoastalBC(A6andA7)ContributionsfromlocalCUswerehigherinsamplesfrommoreinshoreareas(A7-7August212southernCUsandregions)withcontributionsfromsouth-ernCUshigherinthemoreseawardareas(A6-9September862southernCUsandregions)LikethewestcoastofVancouverIslandmigrating stocks were more likely to be found in more seawardlocations
emspensp emsp | emsp11BEACHAM Et Al
33emsp|emspEvaluation of 2017 fishery sampling
Onesourceoffisherysampleswasdirectfisherysamplingofindi-viduals(bothadiposefin-clippedandunclipped)wherethefocuswas not associated with potential CWT recovery In this casesamplingofunclippedindividualsfromthenorthern(AreaF)trollfishery indicated that initially (July 10ndash12) northern and centralBCCUscontributed23tothecatchsouthernBCCUs53andsouthern US regions 23 (Supporting Information Table S3)Stock composition changed markedly in samples collected July13ndash18 with Alaska contributing 12 northern and central BCCUs contributing 72 and southern BCCUs contributing 16withnosouthernUScontributionSamplescollectedduringJuly22ndash28revealedadominantcontributionfromnorthernandcen-tralBCCUs(96)andCUsfromthisregioncontributedbyfarthelargestportionof the catch in subsequent sampleswith south-ernBCCUsandsouthernUSregionslargelyabsent(SupportingInformation Table S3) The average contribution to the weeklysamples from July 22 onwardswas 17 southeast Alaska 16HaidaGwaiindashGrahamIslandLowlandsCU15DouglasChannelndashKitimatArmCU 12Hecate StraitMainlandCU 8NorthernCoastalStreamsCUand7LowerNassCUNotunexpectedlysampling of only clipped individuals from the fishery indicatedverysubstantialdifferencesinstockcompositionbetweenclippedandunclippedindividualsTheaveragecontributiontotheweeklysamples of clipped individuals from July 19 onwards was 32coastalWashington11mid-PugetSound10westVancouverIsland CU and 5 for each of northern Puget Sound and Juande Fuca Strait (Supporting Information Table S4) However theobservedcliprateinthefisherywas16sothecatchwasdomi-natedbyunclippedindividualsTheSeptember2ndash8samplefromthe Haida Gwaii recreational fishery near Langara displayed astockcompositionof42HaidaGwaiindashGrahamIslandLowlandsCU15HecateStraitMainlandCU10LowerSkeenaCUand8 Northern Coastal Streams CU similar to that observed inthe troll fisherywith the local CU (HaidaGwaiindashGraham IslandLowlands) inhigherproportionsaswouldbeexpectedinamoreinshorefishery(SupportingInformationTableS3)
Directsamplingof thecentralcoast troll fisheryrevealedthatonaverage theDouglasChannelndashKitimatArmCUwasthe largestcontributortothefishery(24)followedbytheNorthernCoastalStreamsCU(19)BellaCoolandashDeanRivers(18)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(10) (Supporting InformationTableS3)withallCUsproximaltothelocationofthefisherySamplesfromtherecreationalcentralcoastfisheryindicatedthattheBellaCoolandashDeanRiversCUwasthedominant contributor to the fishery (32) followedby theRiversInletCU(19)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(15)andDouglasChannelndashKitimatArmCU(7) (SupportingInformationTableS3)GiventhelocationofthefisherydominantcontributionsfromtheseCUsweretobeexpected
AsnotedpreviouslysampleswerealsoavailablefromvirtuallyallindividualssampledfromcommercialrecreationalandFirstNationsfisheriesthatwouldberoutinelyprocessedforCWTrecoveryinayearbyacentrallaboratoryIndividualheadsfromcommercialfish-eries sent to the central laboratoryhadbeenpreviously screenedelectronically for thepresenceofaCWTVoluntaryheadsamplesfromadiposefin-clippedindividualsfromrecreationalfisheriesweresenttothecentrallaboratorybutmanyoftheseindividualsdidnotcontainaCWTastherewasnopriorscreeningof these individu-als Stock composition of clipped individuals in the northern trollfishery has been previously outlined above Stock composition inthenorthern recreational fishery indicatedawidegeographicdis-tributionofcontributions rangingfromcoastalWashington (16)thewestcoastofVancouverIslandCU(10)theupperSkeenaCU(9)southeastAlaskaregion(8)mid-PugetSound(7)andeastcoast of Vancouver IslandndashStrait of Georgia CU (6) (SupportingInformation Table S4) Stock composition of clipped individuals inthecentralcoastrecreationalfisherywasdominatedbytheHoweSoundndashBurrardInletCU(27)andeastVancouverIslandndashStraitofGeorgiaCU(25)(SupportingInformationTableS4)butgiventhelowobservedclip rate in the fishery (014) therewasvery littlecatchthatcouldbeidentifiedashatcheryorigin
Directsamplingofrecreationalfisheries insouthernBCwhichincluded both clipped and unclipped individuals in the sample
Population Genotyped Assigned assignment
accuracy of population assignment
Robertson 66 66 1000 1000
Quinsam 69 68 986 1000
Puntledge 21 19 905 1000
BigQualicum 84 84 1000 1000
Goldstream 27 19 704 1000
Chilliwack 195 183 938 1000
Inch 91 85 934 1000
Stave 20 19 950 1000
Total 573 543 948 1000
TA B L E 3 emspPercentageofadiposefin-clippedjacks(age2years)sampledin2016andassignedviaparentageanalysistoparentsin2014broodstockpopulationsPopulationofsamplingwasassumedtobethepopulationoforiginofthejacksforevaluationofaccuracyofassignment
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp11BEACHAM Et Al
33emsp|emspEvaluation of 2017 fishery sampling
Onesourceoffisherysampleswasdirectfisherysamplingofindi-viduals(bothadiposefin-clippedandunclipped)wherethefocuswas not associated with potential CWT recovery In this casesamplingofunclippedindividualsfromthenorthern(AreaF)trollfishery indicated that initially (July 10ndash12) northern and centralBCCUscontributed23tothecatchsouthernBCCUs53andsouthern US regions 23 (Supporting Information Table S3)Stock composition changed markedly in samples collected July13ndash18 with Alaska contributing 12 northern and central BCCUs contributing 72 and southern BCCUs contributing 16withnosouthernUScontributionSamplescollectedduringJuly22ndash28revealedadominantcontributionfromnorthernandcen-tralBCCUs(96)andCUsfromthisregioncontributedbyfarthelargestportionof the catch in subsequent sampleswith south-ernBCCUsandsouthernUSregionslargelyabsent(SupportingInformation Table S3) The average contribution to the weeklysamples from July 22 onwardswas 17 southeast Alaska 16HaidaGwaiindashGrahamIslandLowlandsCU15DouglasChannelndashKitimatArmCU 12Hecate StraitMainlandCU 8NorthernCoastalStreamsCUand7LowerNassCUNotunexpectedlysampling of only clipped individuals from the fishery indicatedverysubstantialdifferencesinstockcompositionbetweenclippedandunclippedindividualsTheaveragecontributiontotheweeklysamples of clipped individuals from July 19 onwards was 32coastalWashington11mid-PugetSound10westVancouverIsland CU and 5 for each of northern Puget Sound and Juande Fuca Strait (Supporting Information Table S4) However theobservedcliprateinthefisherywas16sothecatchwasdomi-natedbyunclippedindividualsTheSeptember2ndash8samplefromthe Haida Gwaii recreational fishery near Langara displayed astockcompositionof42HaidaGwaiindashGrahamIslandLowlandsCU15HecateStraitMainlandCU10LowerSkeenaCUand8 Northern Coastal Streams CU similar to that observed inthe troll fisherywith the local CU (HaidaGwaiindashGraham IslandLowlands) inhigherproportionsaswouldbeexpectedinamoreinshorefishery(SupportingInformationTableS3)
Directsamplingof thecentralcoast troll fisheryrevealedthatonaverage theDouglasChannelndashKitimatArmCUwasthe largestcontributortothefishery(24)followedbytheNorthernCoastalStreamsCU(19)BellaCoolandashDeanRivers(18)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(10) (Supporting InformationTableS3)withallCUsproximaltothelocationofthefisherySamplesfromtherecreationalcentralcoastfisheryindicatedthattheBellaCoolandashDeanRiversCUwasthedominant contributor to the fishery (32) followedby theRiversInletCU(19)SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthernFjordsCU(15)andDouglasChannelndashKitimatArmCU(7) (SupportingInformationTableS3)GiventhelocationofthefisherydominantcontributionsfromtheseCUsweretobeexpected
AsnotedpreviouslysampleswerealsoavailablefromvirtuallyallindividualssampledfromcommercialrecreationalandFirstNationsfisheriesthatwouldberoutinelyprocessedforCWTrecoveryinayearbyacentrallaboratoryIndividualheadsfromcommercialfish-eries sent to the central laboratoryhadbeenpreviously screenedelectronically for thepresenceofaCWTVoluntaryheadsamplesfromadiposefin-clippedindividualsfromrecreationalfisheriesweresenttothecentrallaboratorybutmanyoftheseindividualsdidnotcontainaCWTastherewasnopriorscreeningof these individu-als Stock composition of clipped individuals in the northern trollfishery has been previously outlined above Stock composition inthenorthern recreational fishery indicatedawidegeographicdis-tributionofcontributions rangingfromcoastalWashington (16)thewestcoastofVancouverIslandCU(10)theupperSkeenaCU(9)southeastAlaskaregion(8)mid-PugetSound(7)andeastcoast of Vancouver IslandndashStrait of Georgia CU (6) (SupportingInformation Table S4) Stock composition of clipped individuals inthecentralcoastrecreationalfisherywasdominatedbytheHoweSoundndashBurrardInletCU(27)andeastVancouverIslandndashStraitofGeorgiaCU(25)(SupportingInformationTableS4)butgiventhelowobservedclip rate in the fishery (014) therewasvery littlecatchthatcouldbeidentifiedashatcheryorigin
Directsamplingofrecreationalfisheries insouthernBCwhichincluded both clipped and unclipped individuals in the sample
Population Genotyped Assigned assignment
accuracy of population assignment
Robertson 66 66 1000 1000
Quinsam 69 68 986 1000
Puntledge 21 19 905 1000
BigQualicum 84 84 1000 1000
Goldstream 27 19 704 1000
Chilliwack 195 183 938 1000
Inch 91 85 934 1000
Stave 20 19 950 1000
Total 573 543 948 1000
TA B L E 3 emspPercentageofadiposefin-clippedjacks(age2years)sampledin2016andassignedviaparentageanalysistoparentsin2014broodstockpopulationsPopulationofsamplingwasassumedtobethepopulationoforiginofthejacksforevaluationofaccuracyofassignment
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
12emsp |emsp emspensp BEACHAM Et Al
indicated that therecouldbeverysubstantialdifferences instockcompositionforfisherysamplestakeninthesamemonthandgen-erallocationbetweendirectfisherysamplesandthosederivedfromsamplesofclippedindividualsdeliveredtothecentralprocessinglab-oratoryforpotentialCWTrecoveryForexampleintheJulysamplefromtheJohnstoneStraitcreelsurvey(N=20415cliprate)twoCUs(SouthernCoastalStreamsndashQueenCharlotteStraitndashJohnstoneStraitndashSouthern Fjords HomathkondashKlinaklini Rivers) comprised31ofthesample(SupportingInformationTableS5)IntheAugustsample(N=6218cliprate)thesetwoCUscomprised52ofthesampleSamplesdeliveredtothecentralprocessinglaboratoryforthesametimeperiodsbothdisplayednocontributionofthesetwoCUstothesampleAsnoadiposefinclippingoccursforpopulationsintheseCUscontributionsoftheseCUstofisherieswouldbeun-knownifonlyclippedindividualsweresampledfromthefisheriesSimilarly in theWCVI recreational fishery substantial differencesin stockcompositionwith respect to theUScomponentwereob-servedbetweendirectfisherysamplingandsamplesofclippedin-dividualsdeliveredtothecentralprocessinglaboratoryforpotentialCWTrecoveryForexampleintheJunedirectsample(N=11613clip rate)USpopulationswereestimated tohavecomprised97ofthesamplewhereasintheclipped-onlysample(N=23)theUSpopulationscomprised698ofthesampleSimilarlyinJulydirectsamplingofthecreelcatch(N=25318cliprate)indicatedthatUSpopulationscomprised138ofthesamplebutwereestimatedtocomprise643oftheclippedsample(N=205)submittedforpo-tentialCWTrecoveryInAugustdirectsamplingofthecreelcatch
(N=178 19 clip rate) indicated that US populations comprised186ofthesamplebutwereestimatedtocomprise685oftheclipped sample (N=205) submitted for potential CWT recovery(Supporting Information Table S5) Clearly sampling only adiposefin-clipped individuals from a fishery cannot provide reliable esti-matesofstockcompositionfromtheentirefishery
ThelargemajorityofsamplesfromrecreationalfisheriesthatwasprocessedbythecentrallaboratoryforpotentialCWTrecoverywasderivedfromfisheriesinsouthernBritishColumbiaIntheJohnstoneStraitfisherytheeastVancouverIslandndashStraitofGeorgiaCUcom-prised31of the seasonal sample followedby the LowerFraserCU(17)HoweSoundndashBurrardInletCU(17)andWashingtonre-gionsSkagitRiver(8)andnorthernPugetSound(8)(SupportingInformation Table S4) In the Strait of Georgia fishery the HoweSoundndashBurrard Inlet CU (36) the lower Fraser River CU (32)andtheeastcoastofVancouverIslandndashStraitofGeorgiaCU(12)werethemaincontributors to thecatchwith lessercontributionsfromnorthernPugetSound(7)andSkagitRiver(6)(SupportingInformationTableS4)IntheJuandeFucaStraitfisherythelowerFraserRiverCU(25)wasthedominantcontributorwiththeHoweSoundndashBurrard Inlet CU (9) and east Vancouver IslandndashStrait ofGeorgiaCU(8)alsocontributingtothefisheryContributionsfromWashington regions were also important among them northernPugetSound(17)mid-PugetSound(11)andsouthPugetSound(10)Giventhelocationofthefisheryalargeobservedcontribu-tion (52) ofWashington-origin coho salmonwould be expected(SupportingInformationTableS4) InthewestcoastofVancouver
Population
2016 jacks 2017 CWT
True GSI PBT‐GSI True GSI PBT‐GSI
Quinsam 120 117(14) 119(14) 74 70(11) 72(11)
Puntledge 37 38(11) 37(10) 03 00(01) 03(02)
BigQualicum 147 148(16) 150(17) 53 59(11) 62(11)
Robertson 115 115(14) 115(13) 135 134(14) 135(14)
Inch 155 72(12) 152(12) 185 62(12) 176(12)
Norrish 02 72(13) 06(13) 00 107(14) 03(14)
Coldwater 00 00(00) 00(00) 08 07(03) 08(04)
Salmon 00 00(00) 00(00) 05 04(03) 05(03)
Chilliwack 340 342(20) 341(20) 00 00(00) 00(00)
Stave 35 42(09) 36(09) 00 00(00) 00(00)
Goldstream 47 35(09) 41(09) 00 00(00) 00(00)
Capilano 00 00(00) 00(00) 00 00(01) 00(00)
Nitinat 00 00(01) 00(01) 00 00(00) 00(00)
Conuma 00 00(00) 00(00) 00 00(00) 00(00)
Rosewall 00 00(00) 00(00) 00 03(03) 02(03)
Tenderfoot 00 00(00) 00(00) 00 00(00) 00(00)
Mamquam 00 00(00) 00(00) 00 00(00) 00(00)
Chehalis 00 00(00) 00(00) 00 00(00) 02(00)
Nicomekl 00 00(00) 00(00) 00 00(00) 00(00)
Serpentine 00 00(00) 00(00) 00 00(00) 00(00)
TA B L E 4 emspEstimatedpopulation-specificstockcompositionofthe2016jacksample(N=573)and2017CWTsample(N=620)for20populationswithestimatesderivedfromGSIonlyandcombinedPBTandGSIresultsStandarddeviationoftheestimatesisinparentheses
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp13BEACHAM Et Al
TAB
LE 5
emspTotalnumberofcoded-wiretags(CWTs)observedin2017CanadianfisherysamplesforpopulationswherePBTcouldbeappliednumberofindividualswithCWTssubmittedfor
subsequentgeneticanalysisnumberofindividualscontainingaCWTsubsequentlysuccessfullygenotypednumberofgenotypedindividualswithCWTsassignedviaPBTnumberof
untaggedindividualsassignedviaPBTtotalnumberofindividualsassignedviaPBTthepercentageofgenotypedindividualswithCWTsfromspecificpopulationsassignedviaPBTandthe
subsequentaccuracy()ofassignmentofcoded-wiretaggedindividualsforpopulationswheretheindividualsweregenotypedinthefisherysamplesandinwhichthe2014broodstockwas
genotyped
Popu
latio
nTo
tal C
WTs
Subm
itted
CW
TsCW
T in
divi
dual
s ge
noty
ped
PBT
assi
gnm
ents
C
WT
assi
gned
a
ccur
acy
of
assi
gnm
ent
With
CW
TsN
o CW
TsSu
m o
f PBT
Quinsam
6262
4641
4788
891
100
0
Puntledge
33
22
13
100
010
00
BigQualicum
4039
3331
4071
939
100
0
Robertson
117
9784
81186
267
964
100
0
Inch
117
117
115
102
610
8887
100
0
Coldwater
96
55
38
100
010
00
Salmon
43
33
25
100
010
00
CWTtotal
352
335
288
265
285
550
920
100
0
Nitinat
00
00
2323
Conuma
00
00
1010
Rosewall
00
00
44
Goldstream
00
00
77
Tenderfoot
00
00
4141
Mam
quam
00
00
1313
Capilano
00
00
135
135
Chilliwack
00
00
253
253
Norrish
00
00
9393
Chehalis
00
00
5757
Stave
00
00
2727
Nicomekl
00
00
66
Serpentine
00
00
1111
Total
288
264
965
1230
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
14emsp |emsp emspensp BEACHAM Et Al
Islandfishery therecreationalcatchwasdominatedbyUS-originindividuals with 66 of the seasonal estimate derived from USregions primarily those in Puget Sound and coastalWashingtonwith somecontribution from theColumbiaRiver (5) (SupportingInformation Table S4)Not unexpectedly themainCanadian con-tributortothefisherywasthewestcoastofVancouverIslandCU(9)alongwiththeeastVancouverIslandndashStraitofGeorgiaCU(7)andlowerFraserCU(7)
Sampleswere also available from freshwater fisheries primar-ilyintheFraserRiverdrainageAsamplefromthemainstemFraserRiver recreational fishery (n=154 PBT assignments=146) indi-cated a stock composition of 56 for Inch CreekNorrish Creek23 Chilliwack River 9 Chehalis River and 9 Stave River AsamplefromNicomenSlough(n=57PBTassignments=55)whereboth Norrish Creek and Inch Creek drain and the creek mouthslt1kmfromeachotherdisplayedastockcompositionof53InchCreekand46NorrishCreekoriginAsamplefromtheChilliwackRiver drainage (n=82 PBT assignments=79) displayed a stockcompositionof100ChilliwackRiveroriginOutsideoftheFraserRiverdrainagethreeindividuals(noCWTs)fromtheCapilanoRiverfishery were identified as Capilano River in origin via PBT and
10 individuals from the SquamishRiver fisherywere identified asTenderfootCreek(partofSquamishRiverdrainage)inoriginviaPBTOnVancouver Island two individuals (onewith aCWT) from theBigQualicumRiver fisherywere identified as BigQualicumRiverinoriginviaPBT13individuals(sixwithCWTs)fromtheQuinsamRiverdrainagewere identifiedasQuinsamRiver inoriginviaPBTandeightindividuals(noCWTs)fromtheSomassRiverfisherywereidentifiedasRobertsonCreek(partofSomassRiverdrainage)inor-iginviaPBT
34emsp|emspComparison of CWT and PBT individual identification
Asgenotypeswereavailable from829of the samplesprovidedbythecentrallaboratorythatprocessedsamplesforCWTrecoveryitwaspossibletomakeadirectcomparisonbetweenthequantityofinformationprovidedbyrecoveryofCWTsandidentificationofindividuals via PBT For the seven populationswhereCWTswereappliedandthe2014broodstockgenotypedtherewere352CWTsrecovered from individuals in these seven populations sampledinCanadian fisheriesOf these individuals 335were sampled for
TA B L E 6 emspPBTassignmentsbypopulationforfisheriesinBCduring2017forsamplessenttoacentrallaboratoryforpotentialCWTrecoveryFisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundandAlberniInletsport(j)freshwatersport(k)allfisheries
Population
Fishery
1 2 3 4 5 6 7 8 9 10 11
Quinsam 4 1 6 51 4 6 2 1 13 88
Puntledge 1 2 3
BigQualicum 1 3 2 25 21 12 5 2 71
Robertson 17 15 2 13 1 1 33 177 8 267
Inch 1 4 8 15 3 77 108
Norrish 7 18 6 3 59 93
Coldwater 2 5 1 8
Salmon 1 4 5
Nitinat 4 4 13 2 23
Conuma 2 6 2 10
Rosewall 1 3 4
Goldstream 1 2 2 2 7
Tenderfoot 3 10 8 8 1 10 41a
Mamquam 1 2 3 3 4 13
Capilano 2 29 71 21 8 1 3 135
Chilliwack 1 24 39 54 14 121 253
Chehalis 9 10 18 6 14 57
Stave 4 3 5 1 14 27
Nicomekl 6 6
Serpentine 1 3 5 2 11
Allpopulations 23 24 6 14 187 195 176 99 184 321 1230a
aIncludesoneindividualofunknownmarinecatchregion
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp15BEACHAM Et Al
potential genotyping (individuals sampled in test fisheries werenotincludedinthesamplestobegenotyped)andgenotypeswereobtainedfrom860(288335)oftheinitial individualsprocessed(Table5)PBTassignmentsweremadefor920(265288)ofthegenotyped individuals and PBT assignmentswere 100 accuratewith respect to population of origin and age in comparison withCWTsTherewere285additionalPBTassignmentsmadefortheseseven populations whichwere individuals that had been adiposefinclippedbutwerenot taggedwithaCWTFor thesamplepro-vided335CWTsfromthesevenpopulationswererecoveredand500PBTassignmentsweremadewith49moreindividualidenti-ficationsthroughPBTthanwithCWTsInaddition680PBTassign-mentsweremadefor13populationswherenoCWTswereappliedwith367 (1230PBTvs335CWT)more individual assignmentsmadeforthesamebasesample(Table5)
35emsp|emspApplication of PBT to Canadian fishery samples
PBTwasappliedto identificationof1230individuals inanumberoffisheriesinBCfromwhichCWTscouldpotentiallyberecoveredwiththeintentofcombiningPBTandGSItoprovidehigh-resolutionestimates of stock composition in the samples from these fisher-ies (Table6) Therewere also269additional individuals identifiedviaPBTprimarilyfromdirectsamplingofthecreelcatchinrecrea-tionalfisheriesinsouthernBCwherebothadiposefin-clippedandunclipped individualswere included in the samples (Table 7) TheRobertson Creek and Quinsam River populations were the mostwide-ranging populations observed with individuals from thesepopulations observed in northern commercial troll and recrea-tionalfisheriesthroughtotheJuandeFucaStraitrecreationalfish-ery at the southendofVancouver Island In contrast thereweresome major production populations such as Capilano River andChilliwackRiverwheremarinefisheryrecoveriesweremainly lim-itedtoJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIslandIdentificationofindividualsfromtheColdwaterRiverandSalmonRivertwopopulationsofcon-servationconcernandcurrentlymarkedwithCWTswasrestrictedprimarilytolocalfisheriesintheStraitofGeorgiaandJuandeFucaStraitPopulationsfromtheFraserRiverdrainagedisplayedamorerestricted geographic distribution in fisheries thandid those fromVancouver Island Contributions to fisheries were observed foreverysinglepopulationwherethebroodstockhadbeengenotypedPBTprovided the firstknownoccurrenceof identificationof indi-vidualsoriginating fromthe13non-CWTpopulations inCanadianfisheries
36emsp|emspEstimation of catch of hatchery‐origin populations
Assessment of the impact of fisheries on specific CUs or popula-tions within CUs via genetics requires that accurate high-resolu-tionestimatesofstockcompositionofthecatchareavailableWe
obtainedtheseestimatesofstockcompositionwithrespecttoCU(SupportingInformationTablesS2ndashS5)andpopulationswithinCUs(Supporting InformationTableS6) toestimate thepopulation-spe-cific catch of adipose fin-clipped individuals for the fisheries out-lined inTable8Foreach fishery theclippedcatch thatwaskeptforindividualpopulationswasestimatedviaseasonalstockcompo-sitionestimatesoftheclippedcatch(SupportingInformationTableS6)Hatchery-origin contributions to thekept catch for thegeno-typedpopulationswereestimatedtobethelargestintheStraitofGeorgiarecreationalfisherywiththeCapilanoRiverandChilliwackRiverpopulationscomprising53oftherecreationalcatch(Table8)Hatchery-origincontributionstothekeptcatchwerethelargestforrecreationalfisheriesinJohnstoneStraittheStraitofGeorgiaJuandeFucaStraitandthewestcoastofVancouverIsland
37emsp|emspAssessment of escapement
Non-broodstockescapementsampleswereavailablefrom13popu-lations andPBT assignmentsweremade for 904 (15301692)of the individuals genotyped Stray rateswere estimated at 07(101530) (Table 9) Strays were usually observed between
TA B L E 7 emspPBTassignmentsbypopulationforfisheriesinsouthernBCduring2017bydirectsamplingFisherieswere(a)JohnstoneStraittestgillnet(b)JohnstoneStraitsport(c)StraitofGeorgiasport(d)JuandeFucaStraitsport(e)westcoastVancouverIslandsport(f)allfisheries
Population
Fishery
1 2 3 4 5 6
Quinsam 6 12 8 3 2 31
Puntledge 1 1
BigQualicum 2 2 12 2 18
Robertson 1 2 6 9
Inch 1 9 3 13
Norrish 1 12 3 16
Coldwater 1 1 2
Salmon 1 1
Nitinat 1 1
Conuma 4 4
Rosewall 3 3
Goldstream 1 2 1 1 5
Tenderfoot 1 1 5 1 8
Mamquam 1 3 4
Capilano 1 6 54 4 2 67
Chilliwack 1 1 50 13 1 66
Chehalis 4 9 13
Stave 1 3 1 5
Nicomekl 1 1
Serpentine 1 1
Allpopulations 14 28 168 43 16 269
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
16emsp |emsp emspensp BEACHAM Et Al
geographically proximate populations such as between RosewallCreekandPuntledgeRiveron theeastcoastofVancouver Islandand between Inch Creek and Norrish Creek both draining intoNicomen Slough as part of the Fraser River drainage The largeststraydistanceobservedwasforfourindividualsestimatedtohavestrayedfromtheChehalisRivertotheStaveRiveradistanceofap-proximately40kmintheFraserRiverdrainage
38emsp|emspEstimation of exploitation rate
Estimation of exploitation rates (ER) for populationswas conductedwithCWTswhenavailableandwithgeneticsifescapementestimateswereavailableFortheQuinsamRiverpopulationtheERestimatedviaCWTs(31)andgenetics(28)weresimilar(Table10)Largercontri-butionsofhatchery-origincatchandescapementestimatedviaCWTsthangeneticswerelikelyattributabletoaportionoftheproductionthat
wasmarkedwithCWTsbutwerenotadiposefinclippeduponreleaseForthePuntledgeRiverpopulationwhereonly128ofthejuvenileproductionwasclippedtheERestimatedviageneticswashigher(32)thanviaCWTs(10)(Table10)CatchestimatesforthispopulationviaCWTswerederivedfromtheexpansionofthreerecoveredCWTssoeachrecoveredCWTwasequivalenttoa3ERwithsamplingvari-abilityandCWTlossduringsamplingpotentiallyaccountingformuchoftheobserveddifferenceforthispopulationFortheBigQualicumRiverpopulationbothCWTsandgeneticsprovidedanestimateofERof22Adiscrepancy inERwasobservedfor theRobertsonCreekpopulationwitharateof44fromCWTscomparedwith27fromgenetics(Table10)DifferenceswereapparentinboththeestimatedcatchandhatcherycomponentoftheescapementatRobertsonCreekAtInchCreekinthelowerFraserRivertheERestimatedviaCWTswas41and35viagenetics(Table10)LiketheQuinsamRiverpopula-tionaportionoftheproductionwasmarkedwithCWTsbutwasnot
TA B L E 8 emspCatchofhatchery-origincohosalmonbypopulationforfisheriesinBCduring2017withcatchderivedfromPBT-GSIfor10fisheriesinBCHatchery-origincatchwasestimatedas(totalcatch)(observedadiposefincliprateinthecatch)Population-specifichatchery-origincatchwasestimatedas(hatchery-origincatch)(population-specificstockcomposition)Fisherieswere(a)northerntroll(b)northernsport(c)centraltroll(d)centralsport(e)JohnstoneStraitsport(f)StraitofGeorgiasport(g)JuandeFucaStraitsport(h)westcoastVancouverIslandsportandtroll(i)BarkleySoundndashAlberniInletsport(j)freshwatersport(k)allfisheriesNisthenumberofindividualsincludedintheseasonalsampleforestimatedstockcompositionofthecatch
Population 1 2 3 4 5 6 7 8 9 10 11
Catch 339623 35100 6448 18180 7833 7636 8121 21083 5823
Cliprate() 164 164a 014b 014 4004 8393 8358 7102 2112
Hatchery-origincatch 5570 576 9 27 3136 6409 6788 14974 1230
N 768 85 9 27 192 171 374 517 213
Quinsam 56 6 0 6 558 122 102 75 6 204 1135
Puntledge 0 13 1 0 22 0 0 45 0 0 81
BigQualicum 50 9 2 0 339 596 292 360 0 31 1679
Robertson 596 59 2 0 144 25 14 1078 1069 123 3110
Inch 6 0 0 0 38 192 238 90 0 924 1488
Norrish 0 0 0 0 75 481 122 120 1 768 1567
Coldwater 0 0 0 0 0 45 82 30 0 0 157
Salmon 0 0 0 0 0 26 61 0 0 0 87
Nitinat 156 24 0 2 0 0 61 465 15 0 723
Conuma 22 0 0 0 28 0 0 300 49 0 399
Rosewall 11 0 0 0 9 71 0 60 0 0 151
Goldstream 150 7 0 0 28 0 82 479 0 0 746
Tenderfoot 22 1 1 4 145 333 163 30 0 153 852
Mamquam 23 0 0 2 28 71 68 0 4 0 196
Capilano 78 0 0 2 345 1904 407 300 7 46 3089
Chilliwack 22 0 0 1 245 923 883 435 0 2998 5507
Chehalis 0 0 0 0 107 282 313 180 0 42 924
Stave 11 0 0 0 57 141 102 60 10 42 423
Nicomekl 0 0 0 0 6 0 102 0 0 0 108
Serpentine 6 0 0 0 10 83 81 60 0 0 240
AllPBTpopulations 1209 119 6 17 2184 5295 3173 4167 1161 5329 22662aAdiposefinclipratewasassumedtobeequivalenttothatobservedinthetrollfisherybAdiposefinclipratewasassumedtobeequivalenttothatobservedintherecreationalfishery
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp17BEACHAM Et Al
adiposefinclippeduponreleasepossiblyaccountingforpartofthedif-ferenceinestimatedERsERsfortheSalmonRiverandColdwaterRiverpopulationsestimatedviaCWTswereabout6lessthanthoseesti-matedviageneticswhichwaslargelyattributabletohigherestimatedfisherycatchesofthesepopulationsviageneticsthanwasobtainedviaCWTs(Table10)BothofthesepopulationsoriginatefromCUsofcon-servationconcernandtheERsforthesepopulationswereamongthelowestobservedERsofthesevenindexpopulationsexamined
AnaloguestoERswerealsodeterminedforeightpopulationswherenoCWTswereappliedTheyrangedfrom20fortheConumaRiverpopulationenhancedinalargehatcheryonthewestcoastofVancouverIslandto96fortheGoldstreamRiverpopulationapopulationatthesouthernendofVancouverIslandwhereproductionissupplementedby a small volunteer-staffed hatchery (Table 10) Estimated ERs ofpopulationsfromthelargerproductionhatcherieslikeConumaRiver(20)CapilanoRiver(21)andChilliwackRiver(28)estimateden-tirelyviageneticsweresimilartothoseofotherlargeproductionhatch-eriessuchasBigQualicumRiver (22)RobertsonCreek (27)andQuinsamRiver(28)whereERwasestimatedviaCWTsorgenetics
4emsp |emspDISCUSSION
Canadiancommercialandrecreationalfisheriesforcohosalmonhavebeen severely restricted since the late 1990s but comprehensive
evaluation of the benefits derived from reduced exploitation andmassmarkingofhatcheryproductionhasnotbeenpossibleLargeuntaggedhatcherypopulationsinsouthernBCandlightlymonitoredwildpopulationsofcentralandnorthernBChavebeeninvisibletotheCWT-basedmanagementsystemThesuccessfulapplicationofcurrent genetic technologies in this study allowingaccurate iden-tificationof coho salmon sampled frommixed-stock fisheries hasenabled the first Canadian assessment of fishery impacts that issufficiently informative for conservation-based management asenvisaged in theWSP Coho salmon harvested in Canadian com-mercialandrecreationalfisherieswere identifiedtoCanadianCUsandAmericangeographicregionsfromsoutheastAlaskatoOregonconfirmingtheutilityofaPBT-GSIapproachforconservation-basedassessmentofhatcheryenhancementandmixed-stockharvestonawidegeographicscale
41emsp|emspAccuracy of estimation of stock composition
OnekeydifferencebetweentheCWTmethodandPBT-GSImethodasappliedtosalmonassessmentrelatestotheinabilityoftheCWTapproach to provide estimates of stock composition of the catchCWTrecoveriesareusedtoestimatetotalcontributionsfromtaggedpopulationsthroughldquoexpansionsrdquotoaccountfortheCWTmarkingrateandproportionofthecatchsampledHowevernoestimationof the catch contributions from untagged populations is possible
TA B L E 9 emspNumberofadiposefin-clippedjacksandadultssuccessfullygenotypedfrom2017non-broodstockescapementsamplesthenumberofindividualsofeachlifestagesubsequentlyassignedviaPBTtoeitherthe2014broodstock(adults)or2015broodstock(jacks)numberofindividualsstrayingintoescapementidentifiedviaPBTandestimatedstockcomposition()ofescapementsample(jacksandadultscombined)Standarddeviationofstockcompositionisinparentheses
Population Life stage Number genotyped Assigned via PBT Strays via PBT stock composition
Robertson Jack 10 9 0 1000(10)
Adult 90 90 0
Conuma Adult 14 8 0 934(93)
BigQualicum Jack-Adult 100 100 0 1000(10)
Puntledge Adult(clipped) 4 4 1a 973(34)
Jack-Adult(unclipped) 42 26 0
Quinsam Jack-Adult 196 191 0 983(15)
Capilano Adult 85 68 0 1000(18)
Chilliwack Jack 100 58 0 999(08)
Adult 200 183 0
Inch Jack 557 525 3b 999(03)
Adult 95 90 1b
Norrish Adult 6 4 1c 1000(122)
Stave Adult 111 98 4d 959(21)
Chehalis Adult 75 69 0 1000(13)
Coldwater Adult 6 6 0 1000(125)
Nicomekl Adult 1 1 0 1000(10)
Allpopulations 1692 1530 10 993aRosewallCreekoriginbNorrishCreekorigincInchCreekorigindChehalisRiverorigin
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
18emsp |emsp emspensp BEACHAM Et Al
TAB
LE 1
0emspObservednumberofCWTsfrom2017Canadianfisherysamplingestimated2017catchinCanadianfisheriesofhatchery-originadultcohosalmonviaCWTsER()viaCWTs
ofadultcohosalmoninCanadianfisheriesestimated2017escapementofhatchery-originadultcohosalmonobservedassignmentsviaPBTfrom2017fisherysamplingestimatedcatchvia
PBT-GSIERofclippedadultcohosalmontotaladult2017escapementadiposefincliprateobservedinescapementandclippedhatcherycomponentofadultescapementforselectedcoho
salmonpopulationsNAisnotapplicableasCWTswerenotappliedtothepopulation
Popu
latio
n
Code
d‐w
ire ta
gsPB
T‐G
SIA
dipo
se c
lips
Cat
chEs
cape
men
tC
atch
Esca
pem
ent
Obs
CW
Ts
(adu
lts)
Estim
ated
cat
ch
(adu
lts)
ERH
atch
ery‐
orig
inO
bs P
BTEs
timat
ed c
atch
ERTo
tal a
dults
Clip
rate
Hat
cher
y‐or
igin
clip
ped
Quinsam
621323
313010
881135
283480
823
a2864
Puntledge
314
210
b1330
381
322397
71b
170
BigQualicum
401365
224834
711679
226933
849
5886
Robertson(swim-ins)
c117
4030
445206
267
3110
279511
880
8370
Inch
117
1879
412781
108
1488
352779
997
2771
Coldwater
997
1651
88
157
232178
241
525
Salmon
451
1045
35
8716
676
654
442
Nitinat(swim-ins)
dNA
23723
33e
1497
961
1439
Conuma
NA
10399
20e
3910
419
1638
Rosewall
NA
415
165
f80
100
080
Goldstream
NA
7746
9699
333
33
Tenderfoot
NA
4185
255
f925
747
691
Capilano
NA
135
3089
21e
12244
969
11864
Chilliwack(swim-ins)
NA
253
5507
28e
14492
977
14159
Chehalis(swim-ins)
NA
57924
72f
587
615
361
a CWThatchery-originincludesunclippedtaggedwhichdonotgetcountedinclippedhatchery-originforQuinsamRiverb In2014broodyear128ofPuntledgefryreleasewasclippedandtaggedand
theremainderwasreleasedunclippedIn2017escapementsamplingaCWTlossrateof31wasobservedduetosamplingforotolithsandCWTsfromthesameindividualc TotalescapementtotheStamp
RiverplusRobertsonCreekwas21175adults9511adultsisthehatcheryreturnwhichisallthatwassampledd TotalescapementtotheNitinatRiverwas4883adultsCliprateisfromswim-inssototal
adultsisjustswim-inssameasRobertsonCreeke M
inorcomponentsoftheescapementwerenotenumeratedhowevertheexploitationratecanbeconsideredreasonablyaccurateduetotheveryhigh
percentageoftotalreturnthatwereenumeratedinthehatcheryswim-incountatNitinatRiverCapilanoRiverandChilliwackRiverf Significantcomponentsoftheescapementwerenotenumerated
consequentlyexploitationratewasoverestimatedbyalargebutunknownpercentage
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp19BEACHAM Et Al
precludingtheestimationofstockcompositionfortheentirefisherysamplethatincludesfishfromtaggedanduntaggedpopulationsIncontrastthemainfunctionofaPBT-GSIapproachistoestimatethestockcompositionofthecatchPopulation-specificcatchestimatescanbedeterminedbymultiplyingtheknowncatchbythepopula-tion-specificproportionestimatedviaPBT-GSIIftheproportionofadiposefin-clippedindividuals inthecatchisestimatedandsepa-rateestimatesofstockcompositionforclippedandunclippedindi-vidualsavailablethenpopulation-specificestimatesofcatchofbothclippedandunclippedportionsofthecatchcanbedeterminedIftheproportionofclippedindividualsinthecatchisunknownthenpop-ulation-specific estimates of the clipped (hatchery) and unclipped(natural)portionsofthecatchcanstillbedeterminedifthepropor-tionofclippedindividuals inthepopulationescapement isknownwiththeassumptionthatthesameproportionofclippedindividualsobserved in thepopulationescapementwouldbeobserved in theestimatedpopulationcatchBothCWTsandPBT-GSIcanbeusedtoestimatethecatchofthehatcherycomponentofapopulation(likelyevenawild indexpopulation) for laterapplication inestimationoffisheryexploitationratebutonlythePBT-GSIapproachcanprovidereliableestimatesofstockcompositionoftheunmarkedpopulationsinthefisherysample
GSI provided the foundation for stock composition analysis inthe study and it has been demonstrated to provide reliable esti-matesofcohosalmonstockcompositionforCUsorregionalgroupswithanaverageerrorofle10observedinknown-originsamplesEstimatesforCUsorregionsthatdisplayedhighererrorratescouldlikelybe improvedby increasing thenumberofpopulations in thebaselineusedtorepresenttheCUorregionReliableestimationofsingle-population contributions to fishery samples was enhancedbytheadditionofPBTwith theaverageerror rateobservedwithGSIdecliningbyover50relativetothatobservedwithGSI-PBTInparticularforthesevenpopulationsthateachcomprisedlt5oftheknown-originsamplestheaverageerrorwithGSIwas136whilethatwithGSI-PBTwas016AccuratestockcompositionestimatesofrarepopulationsviaGSIhavebeentraditionallydifficult(ReynoldsampTemplin2004Winansetal2001)butifPBTcanbeappliedinconjunctionwithGSIfortheserarepopulationsthenaccuratees-timatesofstockcompositionscanbeobtainedevenfortheserarepopulationswhentheyoccurinthemixturesampleForexampleinthe2017CWTsamplethreepopulationseachcomprisedle10ofthesamplebutthemaximumerrorforthethreepopulationswhenbothaGSIandPBTapproachwasfollowedwas01Thislevelofresolutionforrarepopulationsinsamplesfrommixed-stocksalmonfisheries will likely be of importance in management of specificfisheries
If 100 of a hatchery broodstock is successfully genotypedthereisanexpectationthat100oftheoffspringfromthebrood-stockshouldbeidentifiableviaPBTInactualpracticeforthetwoknown-originsamplesassignmentratesof916(CWTsample)and948(jacksample)wereachievedFailuretoassignsomeindividualsmaybearesultofincompletesamplingofthehatcherybroodstockfailuretogenotypesuccessfullyallsamplesthatwereprovidedor
genotyping error rates in either someof thebroodstock individu-alsoroffspring thatprecludedassignment to the correctparentsTheoriginof individualsnotassignedwassubsequentlyestimatedviaGSIandthehighlevelsofpopulation-specificaccuracy(ge997)for those populations where it was possible to implement bothPBT and GSI for population-specific stock composition indicatedthatobservedassignmentratesgt91viaPBTwereacceptableforhigh-resolution stock identification analysis Genotypes were ob-tainedsuccessfullyfrom829(2533genotypesfrom3054sam-ples) of the samples providedby the central laboratory employedforpotential recoveryofCWTs from the samplesprocessed fromcommercial recreational and First Nations fisheries Genotypingsuccessimprovedduringthecourseofsampledeliverywithinitialtissuesamples largeandgenotypingsuccessofthemreducedbutsubsequentlysmallertissuesampleswereaffixedtotheWhatmansheetspresumablydryingquickerandpreservingDNAqualitythusresultinginhighergenotypingsuccessAlsothestatusofinitialtis-suequality for thesampleswasuncertainassomesamplesdeliv-eredtothecentrallaboratoryforCWTrecoveryweredeemedtobeunsuitableforDNAsamplingFortissuesamplesdirectlyobtainedfromthefisherygenotypingsuccesswashighforbothcommercial(northern troll fishery marked individuals 989 815 genotypesfrom 824 samples) and recreational fisheries (central coast recre-ationalfishery993542genotypesfrom546samples)
42emsp|emspExploitation rate
ERforapopulationiscatch(catch+escapement)andestimationoffisheryexploitationratesisoneofthekeyoutcomesofassessmentsFortheCWTprogramsamplingofthefisheryandescapementena-blesestimationofthenumberofspecifictagcodesobservedinthetwosetsofsamplesThenumberofobservedtagsisexpandedbythemarkingratetoaccountfortheassociatedhatcheryreleasesnottaggedandagainbythesamplingrate inboththefisheryandes-capementinordertoaccountforhatchery-originindividualsintheunsampledportionofeitherthefisheryorescapementsamples ItisthusmandatorytosamplebothfisheryandescapementsinordertorecoverCWTsWhenmassmarkingofhatcheryfishoccursthemissingadiposefincanbeusedasanindicationthattheindividualmaycontainaCWTbutthelargemajorityoftheclippedindividualswillnotcontainaCWT
Incontrastgenetic-basedassessmentbenefitsfrommassmark-ingofhatcheryproductionparticularlywithregardtoescapementsamplingTheproportionofhatchery-originfishintheescapementcanbedeterminedvisuallyastheproportionofindividualsmissingtheadiposefinwithoutanyfurthersamplingrequiredThebrood-stockandnon-broodstockescapementsamplinginthecurrentproj-ect generally indicated very low rates of straying among sampledpopulations (exceptbetween InchandNorrishcreeks) and thus ifthe escapement abundance is known or estimated the hatcheryportion of the escapement for a population can be estimated viatheobserved clip rateNogenotypingofnon-broodstockescape-ment is required in order to estimate the hatchery component of
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
20emsp |emsp emspensp BEACHAM Et Al
theescapementHoweverifsurvivalofdifferentreleasegroupsisrequiredtobeevaluatedescapementsamplingisrequiredtoassignindividualstoparentsandthereforereleasegroup
Somedifferences in estimatedERs for the seven index popu-lations were observed between those estimated via CWTs andthose estimated via genetics As noted previously some portionofthejuvenileproductionfromsomehatcherieswasreleasedwithCWTsbutwereunclipped resulting in anunderestimationof thecatchofhatchery-origin fish if only the adipose fin clipwasusedto identifyhatchery-origin fish in the catch andescapementTheunclippedbut taggedwithCWTsportionof the releasewouldbeaccountedforthroughtheexpansionsofobservedCWTsandthislikely accounted for the greater abundanceof hatchery-producedcohosalmonineithercatchorescapementestimatedviaCWTsforQuinsamRiverPuntledgeRiverandInchCreekTherewerecasesin which the hatchery component of the escapement was largerwhenestimatedwithadiposefinclipratethanwithexpandedCWTestimatesforBigQualicumRiverandRobertsonCreekpopulationsIntheorysomeclippedhatcheryproductionmaynotbeassociatedwithaCWTandthuswouldnotbe included inexpansionsofob-servedCWTsandinpracticeCWTsmaybemissedduringescape-mentsamplingtherebyunderestimatingthehatcherycomponentEstimated exploitation rates for the two Thompson River indexpopulations (SalmonRiverandColdwaterRiver) representingCUsof conservation concernwere about 6 higher estimated via ge-neticsthanwithCWTsTheestimatedhatcherycomponentoftheescapementwasverysimilarbetweenthe twomethodswith thedifference in ER largely a result of higher fishery catch estimatesvia genetics thanwithCWTsTheobservednumberofCWTs re-covered from fisheries forbothpopulationswasquite limited sixCWTsforColdwaterRiverpopulationifthethreeCWTsrecoveredfromalowerFraserRivertestfisherywereexcludedandfourCWTsfortheSalmonRiverInterestinglyalthoughmostproductionfromthesetwohatcheriesisthoughttobemarkedwithCWTs38(38)oftheColdwaterRiverindividualsidentifiedviaPBTinfisherysam-pleswerenot associatedwith aCWT recovery and40 (25)oftheSalmonRiverindividualsinfisherysampleswerenotassociatedwithaCWTrecoveryTheCWTtag lossratefor2014broodyearjuvenilesduringaretentioncheckwasabout10whichwassim-ilartothetaglossrateobservedinthe2017escapementsampling(clipped individualbutnoassociatedCWTColdwater10[661]Salmon12[651])HoweverCWTtaglosscanoccuruptoeightmonthsaftertagapplicationforthesepopulations(DWillisDFOperscomm)and itmaybethat thehigherfisherycatchesdeter-minedviageneticsmayreflectcontinuingtaglossforthesepopu-lationsThe limitednumberofCWTsrecovered fromfisheries forthesetwopopulationsmayresult inafterexpansionsformarkingrateandfisherysamplingahigherlevelofuncertaintyofthecatchabundancerelativetothatavailablefromgeneticswherebycatchis estimated as clipped fishery abundance stock compositionEstimatedpopulation-specificstockcompositionsofknown-originsamplesviageneticsforthesetwopopulationswerehighlyaccurate(Table4)andaslongasestimatedcatchabundanceofclippedcoho
salmon in fisheries was reliable accurate fishery catch estimatesshouldbeavailableviagenetics
AdditionalindexpopulationsforERestimationcanbedevelopedvia genetics if thehatcherybroodstock is genotyped the juvenileproduction isadiposefinclippedatrelease fisheriesaresurveyedfor stock composition andescapements are sampled for thepro-portionofclippedindividualsCurrentlythreeCWTindexpopula-tionsarelocatedwithinasingleCU(eastVancouverIslandndashGeorgiaStrait)andamorediversegeographicdistributionofindexpopula-tionsmaybedesirableWehaveoutlinedprovisionalresultsforeightadditionalpopulationsbutitmustberecognizedthattheERsmaybemaximumexploitationratesinsomecasesastherewasnocom-prehensive sampling conducted on non-broodstock escapementUnder these circumstances the presence of hatchery-producedindividualsthatreturnedtotheriverratherthantheenhancementfacilitywasunaccountedforintheestimationofenhanced(hatch-ery-produced)abundanceandtheexploitationratewouldbeover-estimatedTousetheselocationsasindexsystemsnon-broodstockescapementsurveyswouldneedtobeconductedinordertoprovideacomprehensiveassessmentofhatchery-originescapementUseofPBT-GSItechnologyprovidesasimplemeanstodevelopadditionalindexpopulationswheresomehatcheryproductionoccursby thesimpleadoptionofbroodstockgenotyping juvenilemassmarkingfisherysamplingandestimationoftheescapementanditscliprate
TheestimatedERof96for theGoldstreamRiverpopulationis of note Goldstream River-origin individuals were identified viaPBTinfourfisheries(Tables6and7)andfivefisheriesviaPBTandGSI(Table8)Estimatedstockcompositionoftheclippedportionofthenorthern troll fisherywas27 (SD =06)yetno individualswereidentifiedviaPBTThisfisherywasestimatedtohavecontrib-utedabout20oftheestimatedcatchofthepopulation(Table8)andweassumedthatareliableestimateofcatchwasobtainedTheestimatedhatcheryabundanceof theescapementwas the lowestofanyofthepopulationssurveyed(Table10) lowerthantypicallyestimated for this populationDrainage escapement estimates forthe prior three yearswere2014 675 2015 208 2016 1838 (PMcCullyGoldstreamhatcheryperscomm)oranaverageof907individuals per year In 2017 fewer than100 individualswere es-timatedintheescapementthehatcheryfailedtoobtainadequatebroodstockandgiventhelowbroodstocknumbersnobroodstocksamplingwas conducted The high ER valuewas concordantwiththepoorescapementobservedin2017andalthoughmaybeover-estimatedtosomedegree likelyreflectedtheregimeexperiencedbythepopulation
InessenceforcohosalmoninsouthernBCinwhichmassmark-ingisappliedandwhichdisplaylowstrayratesamongpopulationsestimationoffisheryexploitationratesrequiresonlygenotypingofindividuals in fishery samples and hatchery broodstocks Forwildpopulationswherethereisafenceorweirinoperationsmallclipscouldbetakenfrompotentialspawnersatthefencetoprovidethebasisforgeneticidentificationoftheiroffspringinfisherysamples(Ford Pearsons amp Murdoch 2015) Some portion of the smoltscouldbecapturedandadiposefin-clippedandsubsequentsampling
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp21BEACHAM Et Al
offin-clippedindividualsinfisherysamplingwouldallowforestima-tionof fishery catch for thewildpopulationThe identificationoffin-clipped individuals in theescapementwouldbepossiblebasedonpreviousjuvenilemarkingTheestimatedescapementabundanceandobservedmarkingrateintheescapementwouldallowdetermi-nationoftheestimatedfin-clippedcomponentoftheescapementWithestimatesofclippedcatchofthewildpopulation infisheriesand abundance of the clipped component in the escapement ex-ploitationrateofthewildpopulationcouldbedetermined
43emsp|emspConservation unit management
ThecurrentCWTsystemofassessmentforcohosalmonprovideslittle information formanagementandassessmentofwildpopula-tionsespecially incentralandnorthernBCwhere themajorityofcoho salmon is commerciallyharvestedTheobservedadipose finclipratein2017fisheriesintheregionwasverylow(014ndash164)and only a portion of the clipped individuals contained CWTs Inexcessof98of thecatchwasunclipped rendering thevalueoftheCWTprogram tobeof very limited value formanagementofdirected fisheries (PKatinicDFOpers comm)ThePBT-GSI ap-proachtofisheryassessmentenablescatchbyCUtobedeterminedforanyfisheryintheprovinceandameanstoimplementtheconser-vationharvestbalancethatcouldbeachievedbymanagingacombi-nationofmixed-stockoceanfisheriesandpotentialin-riverfisheriestargetingonlyhealthyCUs(Priceetal2017)providingsubstantialimprovementtobothCUstatusassessmentasrequiredbytheWSP(DFO2005)andMUfisherymanagementTheuseofPBTtoiden-tifymembersofhatcheryorwild indicatorpopulationsandGSItoidentifyremainingindividualsinthecatchidentifiesthepreviouslyunknowncomponentsoftheharvestwhenassessedwithCWTs
44emsp|emspHatchery management
Ageneticmethodofassessmentenableshatcherybroodstockman-agementandassessmentofhatcheryproductionwitheitherharvestaugmentationorconservationgoalsThedramaticdecline incohosalmon abundance that occurred in British Columbia during the1990sspurredtheimplementationofmassmarkingofhatchery-pro-ducedfish toenablemark-selectiverecreational fisheries inwhichonlyhatchery-producedfishwereharvestedHighlevelsofhatcheryproductionweresuspectedtobeacontributingfactortothepoorsurvivalofwildcohosalmonleadingtoanincreasedawarenessoftheneedtomanagehatcheryproductionandassesshatchery-wildinteractions (Beamish et al 2010) In coho salmonmassmarkingenableshatcherymanagerstoensuretheinclusionofnaturallypro-ducedfishinthebroodstockifdesiredandremovalofhatchery-pro-ducedfishatfencesorweirsinthenaturalenvironmenttocontroltherelativeinfluencesofthenaturalandhatcheryenvironmentonhatchery-supplemented populations in which gene flow betweenthetwospawningenvironmentstakesplace(Mobrandetal2005)Moreovermassmarkingcombinedwithparentageanalysisenablesassessmentofthereproductivesuccessofhatchery-producedfish
that return to spawn in thenaturalenvironment (Abadia-CardosaAndersonPearseampGarza2013Fordetal2015)
Whereas the CWT system is impaired by mass marking ofhatchery fish adipose fin clipping of all hatchery fish combinedwithgenetic identification improveshatchery aswell as fisherymanagementandassessmentGeneticidentificationdoesnotre-quirelethalsamplingprovidesthesexofthesampledindividualandallowsnon-lethalsamplingandreleaseoffishatalllifestagesifrequiredIncontrastrecoveryofCWTsrequireslethalsamplingprecludingthesubsequentreleaseofsampledindividualsandde-termination of the sex in juvenile samples Sampling for geneticanalysisrequiresonlyatissuesample(aslittleasamucousswaborscale)foranalysisWhereasCWTsareoflimiteduseinthestudyof hatchery-wild interactions the non-lethal and simple tissuesamplinghasmadegeneticanalysisofinteractionscommonplacein ecological studies (AshtonCampbellAnders PowellampCain2016DensonBrenkertJenkinsampDarden2012SekinoSaitohYamadaHaraampYamashita2005)Moreover thehatcheryped-igree that can be obtained using PBT enables direct estimationofinbreedingandoutbreedingeffectsinhatcheryproductionandestimationofgeneticparameterssuchasheritability(Berejikianetal2017Kozfkayetal2008)
45emsp|emspUtility of PBT‐GSI for Coastwide Management
ThenecessityofmaintainingaviableCWTsystemforsalmonas-sessmentwasrecognizedundertheoriginalPSTbetweenCanadaand theUnitedStates signed in1985WhereasCWT recoverieswerekeyfactorsinthedevelopmentoftheFRAMforcohosalmonparticularly in thebaseperiodof1986ndash1992 theCWTrecoverysystemwassubsequently impairedbythereleaseofadiposefin-clippedsalmonwithnocorrespondingCWTBy2004thePacificSalmonCommission (PSC) convenedanexpertpanel toexaminelimitationsoftheCWTprogramforbothChinooksalmonandcohosalmonandtoevaluatethecapacityofalternativetechnologiestoprovidedata to improve assessmentof salmonThepanel notedthatPBTwouldprovidetheequivalentofCWTrecoverydatabutthatanempiricaldemonstrationwasneededtovalidatetheoreti-calPBTresultsthatsuggestedbroadfeasibility(PSC2005)Withnolarge-scalePBTapplicationsdevelopedintheinterveningyearsthePSCagaincommissionedin2014anevaluationofthefeasibil-ityandcost-effectivenessofdevelopingacoordinatedcoastwidetag recoverysystemusingPBTstipulating thata transition fromthecoastwideCWTsystemtoaPBTsystemwouldmakerequirethat
1 The PBT system generates at least the same information cur-rently generated from the CWT system via run reconstruction(cohort) analyses of estimated recoveries from individual CWTrelease groups
2 ThePBTsystemwouldhavelong-termannualoperatingcostsnogreaterthanorideallysubstantiallylessthanthoseoftheexist-ingCWTsystem
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
22emsp |emsp emspensp BEACHAM Et Al
3 ThecostofacoastwidePBTsystemwassubstantially less thanthatoftheexistingCWTsystemorthatPBTdeliversadditionalornovelinformationnotprovidedbytheexistingCWTsystemtoinformmanagement of fisheries for coho and Chinook salmon(PSC2015)
Satterthwaiteetal(2015)exploredvariousscenariosunderwhichPBTcouldbeexpandedtoacoastwideapplicationforbothChinooksalmonandcohosalmon
ThePBT-GSImethodofassessmentforBCcohosalmonmeetsthethreecriteriaoutlinedbyPSC(2015)TheaverageannualnumberofCanadian-originCWTsrecoveredinBCcommercialrecreationalandFirstNationscohosalmonfisheriesduring2014ndash2016was391tagswith550CWTsrecoveredin2017from14populationsTheav-erageannualnumberofCWTsrecoveredfromescapementsduring2014ndash2016was 4027 tags with 2636 CWTs recovered in 2017from ten populations Fishery sampling for PBT and GSI in 2017providedgenotypesfor8006individuals1499ofwhichwereas-signedtopopulationandbroodyearwithPBTThegeneticsamplingtherefore provided the required identification for assessment andfisherymanagement for almost three times asmany coho salmonharvestedinBCthandidCWTrecoveryThesePBTfisheryrecover-ieswerespreadover20populationswithassignmentstothelargesthatcherycomprising217ofthetotalassignmentsso itwasnotacaseofasinglehatcherypopulationcomprising thebulkofPBTassignments In fact PBT fishery assignments were observed forallpopulationsgenotypedin2014DirectescapementsamplingforPBT in 2017 provided 1692 genotypes from13 populations andthe6739broodstock individualsgenotypedfor32hatchery2017broodstocksbringtheescapementsamplingtotalto8431individu-alsfor32populationsThePBT-GSIsystemofidentificationgener-atedequivalentinformationrelativetotheCanadianCWTprogramforindexpopulationsofcohosalmoninsouthernBCaswellprovid-ing informationonharvest impacts incurredbyhatcheryandwildpopulationsnotincludedintheCWTindexprogram
WeundertooknocomprehensivecostcomparisonbetweenthePBT-GSIandCWTtechnologiesinthecurrentstudybutaprelimi-narycostanalysiscanbemadeasfollowsApproximately820000CWTswereappliedtooffspringfromthe2014Canadianhatcherybroodstocksandwildescapementatanestimatedcostof$169000(820000$017+$30000Beachametal2008)Therefore thecostperfishfortheCanadian-originCWTsrecovered inCanadianfisheries in2017was$169000550=$307ForPBT-GSIthecostof genotyping the 6061 individuals in the 2014 broodstock sam-plingwas$121220(6061$20Beachametal2008)Thereforethe 1230 individuals from the central CWT recovery laboratoryidentified by PBT in 2017 Canadian fishery samples each cost$99 ($1212201230) If the fish from the direct creel samplingand Johnstone Strait gillnet test fishery are included the cost ofgenetic tagging per PBT recovery declined to $81 per individual($1212201499)comparedwithacostof$307perCWTThePBTmethodof identificationprovidedasubstantialcostadvantagefortaggingperindividualidentifiedin2017fisherysamplescompared
withthatprovidedbytheCWTprogramBeachametal(2018)pro-videdevidencethatforChinooksalmoninBCaPBT-GSIassessmentmethodwassubstantiallycheaperthantheexistingCWTprogramIn this case the cost ofCWT application and recovery and read-ing from fisheryandescapement sampleswascomparedwith thecost of genotyping broodstock fishery and escapement samplesFurtherdetailswereoutlinedbyBeachametal(2018)
Moranetal(2018)notedthatthereisacommonconcernamongsome fisherymanagers and staff that ldquoinvestigation of new tech-nological approaches to provide data for salmon fisherymanage-mentdivertsmoniesthatcanbeusedtomaintaintheexistingCWTprogramrdquo (Pacific Salmon Commission Joint CWT ImplementationTeam2015)IncreasinglyhoweverthedeficienciesprovidedbytheCWT-basedFRAMmodelforbothChinook(Moranetal2018)andcohosalmonare revealedbyapplicationofPBT-GSI technologiesAdditionally many GSI projects are already routinely conductedbecause CWTs do not provide adequate information for fisheriesmanagementdecisionsandassessment(egBeachametal2008Bellingeretal2015Satterthwaiteetal2014)Thecurrentdissat-isfactionofCanadianmanagersofnorthernandcentralcoastalcohosalmonfisheriesandcallsforbetterassessmenttoolsfortheman-agementofmixed-stockandin-riverfisheriesinnorthernBC(Priceet al 2017) alsohighlight theneed for an improvedmanagementregimeAdditionalcostsavingsmayaccruefromimplementationofaPBT-GSImanagementsystemasGSIprojectscurrentlyconductedonanancillarybasistotheCWTprogramaremergedintoroutinefisherysamplingavoidingduplicationofeffortandexpense
ThestrongestbenefitsofaPBT-GSImanagementsystemcomefrom the additional information that it can provide not only forimproved fishery management but also for wild population con-servation and management of enhancement programs CurrentlyfewwildpopulationsaremarkedwithCWTsinBCandtheyareas-sumedtobereliableproxiesforcohosalmonpopulationsoverlargegeographicregionsthatmayencompassmultipleCUsForthefirsttimeanalysisofnorthernandcentralcoastfisheriesinthisstudyen-abledcomprehensivedeterminationof fishery impactsoncentralnorthernriversystemsandtheirconstituentCUsSimilarlycontri-butionstorecreationalfisheriesfromallenhancementprogramsofsouthernBCwere identified for the first timeCoho salmon fromun-CWTedenhancementprogrampopulationsinsouthernBCcon-tributed65of thecatch in southernBC recreational fisheries in2017anduntaggedwildpopulationsfromthemainlandinletsform-ingtheeasternborderofJohnstoneandGeorgiastraitsalsomadesubstantialcontributionsMoreoverstrayratesamonghatchery-in-fluenced coho salmonpopulationswereobserved tobe lowwithimportantexceptionsbeingtheInchCreekandNorrishRiverpop-ulationsrearedatInchCreekHatcheryinwhichreciprocalstrayingwasidentifiedInterestinglyalowerrateofreciprocalstrayingwasidentifiedbetweentheChehalisandStaveRiverpopulationspop-ulationsrearedatdifferenthatcheriesandatdistallocationsinthelowerFraserdrainage
Muchof the increasedutilityof thePBT-GSImethodology re-sultsfromthefactthatcompletebroodstocksamplingensuresclose
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp23BEACHAM Et Al
toa100markrateforjuvenilesThisratecomparesveryfavorablywiththecurrentmarkingrateofapproximately10forCWTsatse-lectedhatcheriesThereducedcostperPBT-taggedfishrecoveredversusCWT-taggedfishstemsfromthehighPBTmarkrateandtheabilityto identifyfishfromallhatcheryfacilitiesenablescompletecatchanalysisthroughtherepresentativesamplingofbothadipose-clippedandunclippedindividuals
46emsp|emspFuture developments
The current study has demonstrated the PBT-GSI capability toidentifyBC-origincohosalmontospecificCanadianhatcheriesandCUs provoking considerationof replacementof the currentCWTsystemforcohosalmonassessmentinBCwithaPBT-GSIbasedap-proachThe304-SNPpanelusedinthecurrentstudytogenotypethe2014cohosalmonbroodstocksandtheirjackreturnsatselectedhatcheries(Beachametal2017)andforgenotypingsamplesfromcohosalmonfisheries in2016and2017hassincebeenupgradedA492-SNPpanel nowexistswith additional loci originating fromresearch conducted in a Genome Canada large-scale applied re-searchprojectandhasbeenusedtogenotypethe2016and2017cohosalmonbroodstocksatanexpandednumberofhatcheries ItisanticipatedthatthisenhancedSNPpanelandtheincreasednum-berof facilities atwhichbroodstock genotypinghasoccurredwillprovideimprovedstockcompositionresultsrelativetothoseofthecurrentstudywhenappliedtocohosalmonfisherysamplesin2019IfCanadaweretoimplementaGSI-PBTmethodofassessmentforcohosalmon inplaceof theCWTprogramthencompleteassess-mentofexploitationratesforCanadianpopulationswouldrequiregenetic analysis of samples from American fisheries if CWTs areretainedastheassessmenttoolforAmericancohosalmonpopula-tionsandfisheriesShouldaGSI-PBTmethodofanalysisbedeemedpracticalforAmericanassessmentpurposesitisconceivablethatacoastwideGSI-PBTassessmentmethodcouldbe implemented forcohosalmonfisheries
5emsp |emspSUMMARY
ThisstudyhasdemonstratedthepotentialforimplementationofacomprehensivePBT-GSImethodologyformanagementandassess-ment of coho salmon in British Columbia that will remedy noteddeficienciesofthecurrentCWT-basedmanagementsystemMostimportantlythegenetictechnologyprovidesanimmediatetoolforidentificationofcohosalmontoCUarequirementforimplementa-tionofmanagementofwildpopulationsasmandatedby theWSPforPacificsalmonandataskthatwouldbeprohibitivelyexpensiveusingCWTsMoreoverthePBT-GSItechnologybenefitsfromthemass marking of hatchery-produced salmon thereby facilitatingimprovedhatcherybroodstockmanagementmonitoringofwild-en-hanced fish interactions and theevaluationofhatchery contribu-tions toharvestTheability to identify readilyhatchery-producedsalmonhasbeenrecognizedasanimperativeformanagingtherisks
andassessingthebenefitsofhatcheryproductionofsalmonidsatthedomesticbilateralandinternationallevels(RuggeroneampIrvine2018)InCanadaextensivecohosalmonconservationandenhance-ment efforts conducted for two decades require comprehensiveevaluationandpossiblemodificationthatcannotbeachievedunderthecurrentmanagementsystemThegeneticmethodologydevel-opedinthisstudyprovidesanopportunityforconservation-basedmanagementofCanadiancohosalmoninwhichtheeconomicben-efitofhatcheryproductioncanbereapedwithouttheimpositionofundueandunknownrisktowildpopulations
DATA ARCHIVING
Multi-locusgenotypes forall sampled jacksaswellas individu-als in fishery and escapement samples are available atDRYADdoi identified as Data from Comparison of coded-wire tag-ging with parentage-based tagging and genetic stock identi-fication in a large-scale coho salmon fisheries application inBritish Columbia Canada Journal Evolutionary Applicationshttpsdoiorg105061dryad6c31bf2 Data files 2016_Coho_Jacks_rubias 2016-17_Coho_Fishery_Samples_rubias2017_Coho_Escapement_rubias
ACKNOWLEDG EMENTS
Averysubstantialeffortwasundertakentoobtainsamplesfromcoho salmon sampled in this study The JO ThomasCWT re-coverylaboratoryprovidedtissuesamplesfrommanyindividualsprocessedfrom2017commercialrecreationalandFirstNationsfisheries in BC as well as samples of adipose fin-clipped indi-vidualsfromthenorthernBCtrollfisheryWealsoacknowledgeFisheriesandOceansCanada(DFO)staffPKatinicwhocollectedsamples of unmarked coho salmon from the northern BritishColumbia troll fishery and the Duncanby Lodge and LangaraLodgestaff for recreational fisherysamples InsouthernBritishColumbia we thank various staff of the Salmon EnhancementProgram of DFO including hatchery managers and staff forsample collection and coordination LKearey P ZetterbergAPereboom BWright EMiyagi and other staff at DFO SouthCoast Area coordinated the direct sampling of coho catch inthe various recreational fisheries in southernBC aswell as theJohnstoneStrait test fisheryLKeareyprovidedqualitycontrolandassuranceofthesamplesDrECAndersonoftheNationalMarineFisheriesServiceSouthwestFisheriesCentergraciouslymodified thecode forRUBIAS toallowtheGSIanalysis topro-ceedJSupernaultassistedintheCOLONYanalysisThisresearchwascarriedoutinconjunctionwithEPIC4(EnhancingProductioninCohoCultureCommunityCatch)aprojectsupportedinpartbyFisheriesandOceansCanadaandthegovernmentofCanadathroughGenomeCanadaGenomeBritishColumbiaandGenomeQuebec Funding support was also previously provided by theNorthernEndowmentFundofthePacificSalmonCommissionforbaselinedevelopment
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
24emsp |emsp emspensp BEACHAM Et Al
ORCID
Terry D Beacham httporcidorg0000-0003-0987-8445
R E FE R E N C E S
Abadia-CardosaAAndersonECPearseDEampGarzaJC(2013)Large-scale parentage analysis reveals reproductive patterns andheritability of spawn timing in a hatchery population of steelhead(Oncorhynchus mykiss)Molecular Ecology224733ndash4746
AndersonEC(2012)Large-scaleparentageinferencewithSNPsAnefficient algorithm for statistical confidence of parent pair alloca-tionsStatistical Applications in Genetics and Molecular Biology11(5)1544ndash6115httpsdoiorg1015151544-61151833
AndersonECampGarza JC (2006)Thepowerof single-nucleotidepolymorphismsforlarge-scaleparentageinferenceGenetics172(4)2567ndash2582httpsdoiorg101534genetics105048074
ArakiHBerejikianBAFordMJampBlouinMS(2008)Fitnessofhatchery-reared salmonids in thewildEvolutionary Applications1342ndash355
AshtonNKCampbellMRAndersPJPowellMSampCainKD(2016) Evaluatingmicrosatellitemarkers for parentage-based tag-gingofhatcheryburbotNorthwest Science90249ndash259httpsdoiorg1039550460900303
BeachamTDWallaceCMacConnachieCJonsenKMcIntoshBCandyJRhellipWithlerRE(2017)Populationandindividualiden-tification of Coho salmon in BC through parentage-based taggingandgeneticstockidentificationAnalternativetocoded-wiretagsCanadian Journal of Fisheries and Aquatic Sciences741391ndash1410
Beacham TDWallace CMacConnachie C Jonsen KMcIntoshB Candy J R ampWithler R E (2018) Population and individualidentification of Chinook Salmon in British Columbia through par-entage-based tagging and genetic stock identification with singlenucleotidepolymorphismsCanadian Journal of Fisheries and Aquatic Sciences751096ndash1105
BeachamTDWintherIJonsenKLWetkloMDengLampCandyJR(2008)Theapplicationofrapidmicrosatellite-basedstockiden-tification tomanagementofaChinooksalmontroll fisheryoff theQueenCharlotte Islands BritishColumbiaNorth American Journal of Fisheries Management 28 849ndash855 httpsdoiorg101577M06-1671
Beamish R J Sweeting RM LangeK LNoakesD J PreikshotDampNevilleCM(2010)EarlymarinesurvivalofcohosalmonintheStraitofGeorgiadeclinestoverylowlevelsMarine and Coastal Fisheries Dynamics Management and Ecosystem Science2424ndash439httpsdoiorg101577C09-0401
BellingerMRBanksMABatesS JCrandallEDGarza JCSylvia G amp Lawson P W (2015) Geo-referenced abundancecalibrated ocean distribution of Chinook Salmon (Oncorhynchus tshawytscha) stocksacross thewest coastofNorthAmericaPLoS One10(7)e0131276
BerejikianBAHardJJTataraCPVanDoornikDMSwansonPampLarsenDA (2017)Rearingstrategiesalterpatternsofsize-selective mortality and heritable size variation in steelhead trout(Oncorhynchus mykiss) Canadian Journal of Fisheries and Aquatic Sciences74273ndash283
BerthelotCBrunetFChalopinDJuanchichABernardMNoeumllBhellipGuiguenY(2014)Therainbowtroutgenomeprovidesnovelinsights into evolution after whole-genome duplication in verte-brates Nature Communications 5 3657 httpsdoiorg101038ncomms4657
DensonMRBrenkertK IV JenkinsWEampDardenT L (2012)Assessing red drum juvenile stocking in a South Carolina estu-aryusinggenetic identificationNorth American Journal of Fisheries
Management32 32ndash43httpsdoiorg101080027559472011649577
Fisheries and Oceans Canada (2005) Canadarsquos policy for conservation of wild Pacific salmon (p57)VancouverBCFisheriesandOceansCanada
FlaggTA(2015)BalancingconservationandharvestobjectivesAre-viewofconsiderationsforthemanagementofsalmonhatcheriesintheUSPacificnorthwestNorth American Journal of Aquaculture77367ndash376httpsdoiorg1010801522205520151044058
FordMPearsonsTNampMurdochA(2015)ThespawningsuccessofearlymaturingresidenthatcheryChinookSalmoninanaturalriversystemTransactions of the American Fisheries Society144539ndash548httpsdoiorg1010800002848720151009561
Hatchery Scientific Review Group (HSRG) (2014) On the science of hatcheries an updated perspective on the role of hatcheries in salmon and steelhead management in the Pacific Northwest Retrieved fromhttpshatcheryreformus
Hess J E AckermanMW Fryer J KHasselmanD J Steele CAStephensonJJhellipNarumSR(2016)Differentialadultmigra-tion-timingandstock-specificabundanceofsteelheadinmixedstockassemblagesICES Journal of Marine Science732606ndash2615httpsdoiorg101093icesjmsfsw138
HilbornR(1992)HatcheriesandthefutureofsalmonintheNorthwestFisheries 17 5ndash8 httpsdoiorg1015771548-8446(1992)017aabbb0005HATFOSaaabb20CO2
Hinrichsen R A Steele C A Ackerman M W Campbell M RNarumSRHessMAhellipMaschhoffBL(2016)Maximumlikeli-hoodestimationoftheproportionofhatchery-originfishonspawn-inggroundsusingcodedwiretaggingandparentage-basedtaggingTransactions of the American Fisheries Society145671ndash686httpsdoiorg1010800002848720161146163
HoltCAampIrvineJR(2013)DistinguishingbenchmarksofbiologicalstatusfrommanagementreferencepointsAcasestudyonPacificSalmoninCanadaEnvironmental Conservation40345ndash355httpsdoiorg101017S0376892913000209
HoltbyBLampCirunaKA(2007)Conservation units for Pacific salmon under the Wild Salmon PolicyCanadianScienceAdvisorySecretariatScienceAdvisoryReport2007070
IrvineJRampFraserGA(2008)CanadarsquosWildSalmonPolicyandthemaintenanceofdiversityAmerican Fisheries Society Symposium49391ndash398
JeffertsKBBergmannPKampFiscusHF(1963)Acodedwiretagidentification system for macro-organisms Nature (London) 198460ndash462
JonesKKCornwellTJBottomDLSteinSampAnlauf-DunnKJ(2018)Populationviability improvesfollowingterminationofcohosalmonreleasesNorth American Journal of Fisheries Management3839ndash55
Jones O W amp Wang J (2010) COLONY A program for par-entage and sibship inference from multilocus genotypedata Molecular Ecology Research 10 551ndash555 httpsdoiorg101111j1755-0998200902787x
Kozfkay C C Campbell M R Heindel J A Baker D J Kline PPowellMSampFlaggT(2008)AgeneticevaluationofrelatednessforbroodstockmanagementofcaptiveendangeredsockeyesalmonOncorhynchus nerka Conservation Genetics91421ndash1430
McClureMMUtterFMBaldwinCCarmichaelRWHassemerPFHowellPJhellipPetroskyCE(2008)EvolutionaryeffectsofalternativeartificialpropagationprogramsImplicationsforviabilityof endangered anadromous salmonidsEvolutionary Applications1356ndash375
MobrandLEBarrJBlankenshipLCamptonDEEvelynTPTFlaggTAhellipSmokerWW(2005)HatcheryreforminWashingtonstateFisheries3011ndash23httpsdoiorg1015771548-8446(2005)30[11HRIWS]20CO2
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711
emspensp emsp | emsp25BEACHAM Et Al
Moran B M amp Anderson E C (2018) Bayesian inference fromthe conditional genetic stock identification model Canadian Journal of Fisheries and Aquatic Sciences httpsdoiorg101139cjfas-2018-0016
MoranPDazeyJLaVoyLampYoungS(2018)Geneticmixtureanal-ysis supports recalibration of the fishery regulation assessmentmodel Fisheries4383ndash97httpsdoiorg101002fsh10017
PacificFisheryManagementCouncil (PFMC) (2008)Fishery Regulation Assessment Model (FRAM) ndash An overview for Chinook and Coho ndash v30 PortlandORPacificFisheryManagementCouncil
Pacific Salmon Commission (PSC) (2005) Expert Panel on the Future of the Coded Wire Tag Program for Pacific Salmon Pacific SalmonCommission Technical Report Number 18 230 p Retrievedfrom httpswwwpscorgpublicationsworkshop-reportscoded-wire-tag-program-review
Pacific Salmon Commission (PSC) (2015) CSC complete PBT review ReviewofSatterthwaiteetal2015Multidisciplinaryevaluationofthe feasibility of parentage-based genetic tagging (PBT) for man-agement of Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Pacific Salmon Commission Joint CWT Implementation Team (2015)Five‐year synthesis report of the PSC Coded Wire Tag (CWT) Improvement ProgramPacificSalmonCommissionTechnicalReportNumber33VancouverBritishColumbia
PriceM H H English K K Rosenberger A GMacDuffeeM ampReynolds J D (2017) Canadarsquos Wild Salmon Policy An assess-ment of conservation progress in British Columbia Canadian Journal of Fisheries and Aquatic Sciences741507ndash1518httpsdoiorg101139cjfas-2017-0127
PriceM H H Rosenberger A G J Taylor G G amp Stanford J A(2014)CommentonldquoPopulationstructureandruntimingofsockeyesalmonintheSkeenaRiverBCrdquoNorth American Journal of Fisheries Management341167ndash1170
Regional Mark Processing Center (RMPC) (2018) RMIS database Retrievedfromhttpswwwrmpcorghomehtml
ReisenbichlerRRampHartmannNA Jr (1980)Effectofnumberofmarkedfishandyearsofrepetitiononprecisioninstudiesofcontri-butiontoafisheryCanadian Journal of Fisheries and Aquatic Sciences37576ndash582httpsdoiorg101139f80-073
ReynoldsJHampTemplinWD(2004)DetectingspecificpopulationsinmixturesEnvironmental Biology of Fishes69233ndash243httpsdoiorg101023BEBFI000002287738588f1
RuggeroneGTampIrvineJR(2018)Numbersandbiomassofnatural-andhatchery-originpinksalmonchumsalmonandsockeyesalmoninthenorthPacificOcean1925ndash2015Marine and Coastal Fisheries Dynamics Management and Ecosystem Science10152ndash168httpsdoiorg101002mcf210023
Sandercock F K (1991) Life history of coho salmon (Oncorhynchus kisutch)InCGrootampLMargolis(Eds)Life history of Pacific Salmon (pp395ndash406)VancouverBCUniversityofBritishColumbiaPress
SatterthwaiteWAnderson E CampbellMGarza J CMohrMNarumSampSpeirC (2015)Multidisciplinary evaluation of the fea‐sibility of parentage‐based genetic tagging (PBT) for management of
Pacific salmon Retrieved from httpswwwpscorgpublicationsworkshop-reportspsc-special-reports
Satterthwaite W H Mohr M S OrsquoFarrell M R Anderson E CBanksMA Bates S JhellipPalmer-ZwahlenM L (2014)Use ofgeneticstockidentificationdataforcomparisonoftheoceanspatialdistributionsizeatageandfisheryexposureofanuntaggedstockand its indicator California coastal versus Klamath River ChinookSalmonTransactions of the American Fisheries Society143117ndash133httpsdoiorg101080000284872013837096
Sekino M Saitoh K Yamada T Hara M amp Yamashita Y (2005)Genetic tagging of released Japanese flounder (Paralichthys oliva‐ceus)basedonpolymorphicDNAmarkersAquaculture24449ndash61httpsdoiorg101016jaquaculture200411006
StarksHAClementoAJampGarzaJC(2016)Discoveryandchar-acterization of single nucleotide polymorphisms in coho salmonOncorhynchus kisutch Molecular Ecology Resources16277ndash287
SteeleCAAndersonECAckermanMWHessMACampbellNRNarumSRampCampbellMR(2013)Avalidationofparent-age-basedtaggingusinghatcherysteelheadintheSnakeRiverbasinCanadian Journal of Fisheries and Aquatic Sciences70 1046ndash1054httpsdoiorg101139cjfas-2012-0451
TompkinsAHamiltonSBatemanJampIrvineJR(2016)Canadian salmon catch and enhanced salmon production in 2014 and 2015 with a historical overview of recreational steelhead catches North PacificAnadromousFishCommissionDocument1654(Rev1)9pFisheriesandOceansCanadaRetrievedfromhttpwwwnpafcorg
Wang J (2016) Individual identification from genetic markerdata Developments and accuracy comparisons of meth-ods Molecular Ecology Research 16 163ndash175 httpsdoiorg1011111755-099812452
WinansG VieleDGrover A Palmer-ZwahlenM TeelD ampVanDoornik D M (2001) An update of genetic stock identifica-tion of chinook salmon in the Pacific Northwest Test fisheriesin California Reviews in Fisheries Science 9 213ndash237 httpsdoiorg10108020016491101753
SUPPORTING INFORMATION
Additional supporting information may be found online in theSupportingInformationsectionattheendofthearticle
How to cite this articleBeachamTDWallaceCJonsenKetalComparisonofcoded-wiretaggingwithparentage-basedtaggingandgeneticstockidentificationinalarge-scalecohosalmonfisheriesapplicationinBritishColumbiaCanadaEvol Appl 2018001ndash25 httpsdoiorg101111eva12711