Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System...

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Evolutionary Applications. 2018;1–25. | 1 wileyonlinelibrary.com/journal/eva Received: 14 June 2018 | Revised: 1 August 2018 | Accepted: 7 August 2018 DOI: 10.1111/eva.12711 ORIGINAL ARTICLE 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. Beacham 1 | Colin Wallace 1 | Kim Jonsen 1 | Brenda McIntosh 1 | John R. Candy 1 | David Willis 2 | Cheryl Lynch 2 | Jean‐Sébastien Moore 3 | Louis Bernatchez 3 | Ruth E. Withler 1 This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. © 2018 The Authors. Evolutionary Applications published by John Wiley & Sons Ltd 1 Fisheries and Oceans Canada, Pacific Biological Station, Nanaimo, BC, Canada 2 Fisheries and Oceans Canada, Regional Headquarters, Vancouver, BC, Canada 3 Département de Biologie, Institut de Biologie Intégrative et des Systèmes (IBIS), Université Laval, Québec, QC, Canada Correspondence Terry D. Beacham, Fisheries and Oceans Canada, Pacific Biological Station, Nanaimo, BC, Canada. Email: [email protected] Funding information Fisheries and Oceans Canada; Genome British Columbia; Pacific Salmon Foundation, Grant/Award Number: EPIC4; Genome Canada Abstract Wild Pacific salmon, including Coho salmon Onchorynchus kisutch, have been sup- plemented with hatchery propagation for over 50 years in support of increased ocean harvest and conservation of threatened populations. In Canada, the Wild Salmon Policy for Pacific salmon was established with the goal of maintaining and restoring healthy and diverse Pacific salmon populations, making conservation of wild salmon and their habitats the highest priority for resource management deci- sion-making. A new approach to the assessment and management of wild coho salmon, and the associated hatchery production and fishery management is needed. Implementation of parentage-based tagging (PBT) may overcome problems associ- ated with coded-wire tag-based (CWT) assessment and management of coho salmon fisheries, providing at a minimum information equivalent to that derived from the CWT program. PBT and genetic stock identification (GSI) were used to identify coho salmon sampled in fisheries (8,006 individuals) and escapements (1,692 individuals) in British Columbia to specific conservation units (CU), populations, and broodyears. Individuals were genotyped at 304 single nucleotide polymorphisms (SNPs) via direct sequencing of amplicons. Very high accuracy of assignment to population (100%) via PBT for 543 jack (age 2) assigned to correct age and collection location and 265 coded-wire tag (CWT, age 3) coho salmon assigned to correct age and release loca- tion was observed, with a 40,774—individual, 267—population baseline available for assignment. Coho salmon from un-CWTed enhanced populations contributed 65% of the catch in southern recreational fisheries in 2017. Application of a PBT-GSI system of identification to individuals in 2017 fisheries and escapements provided high-res- olution estimates of stock composition, catch, and exploitation rate by CU or popula- tion, providing an alternate and more effective method in the assessment and management of Canadian-origin coho salmon relative to CWTs, and an opportunity for a genetic-based system to replace the current CWT system for coho salmon assessment.

Transcript of Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System...

Page 1: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

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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

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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

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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

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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

Page 2: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 3: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 4: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 5: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 6: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 7: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 8: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 9: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 10: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 11: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 12: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 13: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 14: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 15: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 16: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 17: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 18: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 19: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 20: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 21: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 22: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 23: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 24: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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

Page 25: Comparison of coded‐wire tagging with parentage‐based ...456) Beacham...Mark Information System (RMPC, 2018), it was determined that there were 2,664 CWTs (average of 888 CWTs

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