This article was downloaded by: [Oregon State University] On: 19 September 2011, At: 16:22 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Fisheries Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/ufsh20 Development of a Standardized DNA Database for Chinook Salmon L. W. Seeb, A. Antonovich, M. A. Banks, T. D. Beacham, M. R. Bellinger, S. M. Blankenship, M. R. Campbell, N. A. Decovich, J. C. Garza, C. M. Guthrie III, T. A. Lundrigan, P. Moran, S. R. Narum, J. J. Stephenson, K. J. Supernault, D. J. Teel, W. D. Templin, J. K. Wenburg, S. F. Young & C. T. Smith Available online: 09 Jan 2011 To cite this article: L. W. Seeb, A. Antonovich, M. A. Banks, T. D. Beacham, M. R. Bellinger, S. M. Blankenship, M. R. Campbell, N. A. Decovich, J. C. Garza, C. M. Guthrie III, T. A. Lundrigan, P. Moran, S. R. Narum, J. J. Stephenson, K. J. Supernault, D. J. Teel, W. D. Templin, J. K. Wenburg, S. F. Young & C. T. Smith (2007): Development of a Standardized DNA Database for Chinook Salmon, Fisheries, 32:11, 540-552 To link to this article: http://dx.doi.org/10.1577/1548-8446(2007)32[540:DOASDD]2.0.CO;2 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan, sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. FEATURE: GENETICS Developmentof a Standardized DNA Database for Chinook Salmon L. W. Seeb, A.Antonovich, '•qlr • Downloaded by [Oregon State University] at 16:22 19 September 2011 M. A. Banks, T. D. Beacham, M. R. Bellinger, • • S.M. Blankenship, i L M. R. Campbell, N. A. Decovich, J. C. Garza, C.M. Guthrie III, T. A. Lundrigan, • P. Moran, S. R. Narum, J. J. Stephenson, K. J. Supernault, D. J. Teel, W. D. Templin, J.K.Wenburg, '" S. E Young,and C. T. Smith Seebisresearch professor at theSchoolofAquaticandFisheries Sciences, University of Washington, Seattle.Shecan becontacted at Iseeb@u.washington.edu. Antonovichisa biometrician, Decovichisa fishery biologist, andTemplinis a fisheries geneticist at theAlaskaDepartment ofFishandGame,DivisionofCommercial Fisheries, Anchorage. Banks ismarinefisheries geneticist andassociate professor andBellingerisa molecular geneticist at OregonStateUniversity's HatfieldMarineScience Center,Newport.Beacham isa research scientist andSupernault isa research technicianat the Department ofFisheries andOceans Pacific Biological Stationin Nanaimo, BritishColumbia. Blankenship wasformerly with the National Marine FisheriesServiceSouthwestFisheriesScienceCenter in SantaCruz, California, and is now a geneticist attheWashington Department ofFishandWildlife,Olympia. Campbell issenior fisheries research biologist atthe IdahoDepartment ofFishandGame,EagleFishGenetics Laboratory, Eagle.Garzaissupervisory research biologist, National MarineFisheries ServiceSouthwest Fisheries ScienceCenter,SantaCruz,California.Guthrieisa fisheryresearch biologist at the NationalMarineFisheries ServiceAuke BayLaboratories, TedStevensMarineResearch Institute,Juneau,Alaska. Lundrigan wasformerly withtheUniversity ofWashington SchoolofAquaticandFishery Sciences, Seattle,andiscurrently a fisheries biologist at theUniversity ofVictoria's CentreforBiomedical Research, Victoria,BritishColumbia. Moranis a research geneticist andTeelissupervisory fishery biologist at theNationalMarineFisheries Service Northwest Fisheries Science Center,Seattle, Washington. Narumisleadgeneticist andStephenson islaboratory manager at theColumbia River Inter-Tribal FishCommission, Hagerman FishCultureExperiment Station,Hagerman, Idaho.Wenburg isdirectorofgenetics at the U.S. FishandWildlifeServicein Anchorage, Alaska.Youngisa fishandwildliferesearch scientistat the Washington Department ofFishandWildlife,Olympia.Smithwasformerly withtheAlaskaDepartment ofFishandGameDivisionof Commercial Fisheries andisnowa fisheries geneticist at theU.S. FishandWildlifeService,AbernathyFishTechnology Center,Longview,Washington. 540 Fisheries ß vol 32 No 11 ß NOVEMBER 2007 ß WWW.FISHERIES.ORG ABSTRACT: An mternauonal mulu-laboratory projectwasconducted to developa standardized DNA database forChinook salmon(Oncorhynchus tshawytscha). Thisprojectwasin response to theneedsof theChinookTechnicalCommitteeofthePacific SalmonCommission to identifystockcomposition of Chinooksalmoncaughtin fisheries duringtheiroceanicmigrations. Nine genetics laboratories identified13microsatellite locithatcouldbereproducibly assayed in eachof the laboratories. To testthat the lociwerereproducible amonglaboratories, blindtestswereconducted to verifyscoring consistency forthenearly500total alleles.Oncestandardized, a datasetof over16,000Chinooksaltnonrepresenting 110putativepopulations wasconstructed rangingthroughout the areaof interestof the PacificSalmonCommission fromSoutheast Alaskato the Sacramento River •n California.The datasetdifferentiates the majorknowngeneticlineages of Chinooksaltnonandprovides a toolfor genetic stockidentification of samples collected frommixedfisheries. A diverse groupof scientists representing thedisciplines of fishery management, genetics, fishery administration, population dynamics, andsampling theoryarenowdeveloping recommendations forthe integration of thesegeneticdataintooceansalmonmanagement. Desarrollo de una base de datos estandarizada Downloaded by [Oregon State University] at 16:22 19 September 2011 de DNA parael salm6nrey RESUMEN: Se realiz6un proyecto internacional conla participaci6n dediversos laboratorios conla finalidaddedesarrollar unabasededatosestandarizada deDNA parael salm6nrey(Oncorhynchus tshawytscha). Dichoproyecto surgi6comorespuesta a lasnecesidades delComit6T6cnicoChinookdelaComisi6n delSalm6ndelPac•fico paraidentificar lacomposici6n poblacional del salm6nreyqueescapturado pot la pesquerfa durantesumigraci6n. Un totalde nuevelaboratorios de anfilisis gen•ticos tdentificaron y reprodujeron cadauno 13 loci microsat61ites. Con el objetode probarquedichosloci fueranreproducibles entrelaboratorios, secondujeron pruebas an6nimas paraverificarla consistencia de casi500 alelos.Una vezestandarizada, se construy6 unabasededatos construida coninformaci6n proveniente demilsde16,000salmones querepresentan 110poblaciones putativas distribuidas a lo largodel fireade inter6sde la Comisi6ndelSalm6ndel Pacffico, delsureste deAlaskahastael Rfo Sacramento, California.La basededatossirvetantoparaidentificar gen6ticamente losdistintos stocks de salm6nreya partit demuestras combinadas provenientes dela pesquer•a comoparadiferenciar el linajegen6tico conocido m•simportante deesta especie. En la actualidad, un importante grupodecientfficos especializados en disciplinas comoel manejoy administraci6n de pesquerfas, gen6tica,dinfimicapoblacional y teor•adel muestreo estrindesarrollando recomendaciones paraqueestabasede datosgen6ticos seincorpore en el manejodelsalm6n. INTRODUCTION The modelusescatch,escapement, coded- fromthe indicatorstockassigned to them wiretag(CWT) recovery, andrecruitment in the model. The PacificSaltnonTreatywasratified information to forecast relative abundance In 2004,the PacificSalmonCommission in treatyfisheries. conveneda panelof expertsto examine 1985,renegotiated in 1999,and extended With the increaseddependence on limitationsof the CWT prograinin the to the YukonRiver in 2002.Throughthe CWT recoverydata, concernhas been contextof massmarkingand mark-selectreaty,the twonationsagreedto cooper- raisedregardingthe qualityof the CWT tive fisheries,as well as to evaluate the ate in the management,research,and data and inferences drown from those capacityof alternativetechnologies to enhancement of Pacific salmon. Pacific data (Hankin et al. 2005). Historically improveassessment of Chinooksahnon salmonmigratelongdistances duringtheir onlyfishcarrying a CWT hadan adipose (Hankin et al. 2005). The expertpanel marineperiodand are routinelyinter. clip. However,with the adventof mass- concludedthat alternativetechnologies ceptedin fisheries beyondthejurisdiction markingof largenumbersof hatcheryfish couldnot by themselves replaceCWTs, of the government in whosewatersthey usingan adipose clip only (e.g.,Mobrand but thatgeneticstockidentification (GSI) spawn. The PacificSaltnonTreaty,through et al. 2005), recoveryof CWT fishhas could indeedcomplementthe existing the PacificSahnonCommission(PSC), beencomplicated. Recovery nowrequires CWT programs or be usedin combinaservesas a meansto coordinatemanage. both handlingmuch largernumbersof tion with othertechniques suchasotolith ment of the salmon resourceand conduct individuals pairedwith electronicscan- thermalmarkingto estimatethe stock conservation actionsasrequired. ning of thoseadipose-clipped individu- composition of a landedcatchin a parChinooksalmon(Oncorhynchus tshaw- als to detecttags.As a furtherconcern, ticulartime/areafishery(Findings11-13) ytscha)are harvestedthroughoutthe abundance and harvest information is However,they notedlimitationsin the yearby commercial and sportfishersin not available for most Chinook saltnon GSI methods due to the lack of coastwide the waters of southeastAlaska, British stocks,soindicatorstocksare usedwithin geneticbaselines, which are databases of Columbia, and the Pacific Northwest. the modelto represent bothlargergroups genotypes frombreeding populations. Fisheries typicallyharvesthighly mixed of hatchery andwildstocks. The abilityof stocks of Chinook saltnon and are thereFinding 13. ModernGSI methods thehatcheryindicatorstocks to accurately can be used to estimatethe stockcomfore underthe jurisdictionof the Pacific represent wild stockshasbeengenerally position of thelanded catchin a parSaltnonTreaty.Quotasspecified by the supportedin coho saltnon(O. kisutch; ticular time/areafishery. However, PSC aredependent on the abundance of e.g.,Weitkampand Neely 2002),but is the accuracyand precision of data Chinooksaltnon projected byitsChinook still largelyunknownfor manystocksof TechnicalCommittee(CTC) usingthe Chinooksahnon.The wild stocksmay required to estimate stock-age-fishery Chinooksaltnon model(e.g.,CTC 2005). differin ancestry, abundance, andtiming specificexploitation ratesusingGSI between the United States and Canada in Fisherles * vot 32 .o II * NOVEMBER 2007 e VV•/VV. FISHERIES,ORG 541 methods •s dependent upona variety of factors.For example, microsatellite DNA-basedGSI technology isnotyet capable of generating consistent, replicableestimates dueto thelackof a coastwide geneticbaseline, thehistory of stocktransfers withinand among watersheds, and differences in methodologies andmixtureseparation algo- this databasewas comprehensive and vanousmeeungsof salmomdgenet•c•sts usedby multiplelaboratories, a number from 1999to 2001,but progress waslimof limitations,includingthe requirements ited (LaHoodet al. 2002).Finally,a workfor lethalsampling, cryopreservation, lack shophostedby the CTC in 2003 resulted of laboratoryautomation,and the finite in funding for the databasedescribed numberof loci, led researchers to the deciherein.In thefollowingyear,a symposium sionto replacethe allozymebaselinewith on geneticdatabases for fisherymanagea DNA database. rithms.(Hankin et al. 2005) 2004 AmericanFisheriesSociety(AFS) Annual Meeting, and groupsstudying a numberof divergentfisheryresources ableforresolving Chinooksalmonpopula- beganstandardization efforts(e.g.,Atlanuc tion structure.However,unlikeallozymes herring;Marianiet al. 2005).So although where a systemwas developedfor sharthe advantagesof DNA-based markers, ing and transferring data amonglaboraandmicrosatellites in particular, werewell tories, microsatellite baselines evolved documented beginning in the early 1990s for usewithin singlelaboratories (Moran (e.g., Wright and Bentzen 1994; Wirg•n et al. 2006). The largenumberof available microsatellite loci resulted in little andWaldman1994),coordination among laggedmarkedly. overlapamongresearchers' datasets.For laboratories example,at the beginning of 2003over60 The presentdatabase wasconstructed loci were in usefor Chinook salmon,but and evaluatedthrougha two-yearcollabmost(43) wereusedin onlya singlelabo- orative effort of an international muluratory(Figurela). Thiswasnot limitedto agencywork group. Guiding principles Markers basedon DNA, suchas mito- ment and conservation was held at the chondrialDNA (Cronin et al. 1993) and microsatellites (Bankset al. 2000;seeBox Herewereviewa largemulti-laboratory 1), were alsoin useand shownto be valu- effortto developand standardize a replicable DNA databasefor Chinook salmon. Historyand needfor Downloaded by [Oregon State University] at 16:22 19 September 2011 coast.wide databases Beginning in the 1980s, considerable effort and coordination was directed towardsdeveloping standardized allozyme (protein) baselines(e.g., Shaklee and Phelps1990; White and Shaklee1991; seeBox 1). Standardization amonglaboratoriesinvolvesidentifyingandadopting a commonsuiteof loci (specificpolymorphicDNA segments ortheirproducts) and Chinook salmon,butwasthe normfor fishFurthermore, adopting consistent namesfor the various eriesstudiesof manyspecies. even when identical loci were beingused, alleles(seeBox 1). Geneticanalyses using differences in chemistries and instrument allozymedatabases were usedextensively platformsamong laboratoriesproduced to estimate the stock contribution of requiredthat the database would:(1) be subjectto review by scientistsfrom all interestedagencies, (2) be freelyavailable to all researchers managingor studying variable size calls for the same allele from Chinooksalmon,and (3) coverthe range Chinook salmon fisheries in the Columbia the same individual, which precluded easof Chinooksalmonat a geographic scale River, coastalWashington,and Strait of ily merging datasets from multiple regions Juande Fuca(e.g.,Marshallet al. 1991; appropriate to the management objectives Shakleeet al. 1999). Collaborativework (Figurelb). As a result,despitecollection of the PSC with the understanding that duringthe 1990sby multiple state,pro- of a substantial amount of microsatellite the database couldbe easilyexpandedto et vincial,andfederalagencies wasdirected data(e.g.,Nelsonet al. 2001;Beacham includethe entirerangeof the species. It in replacingthe allotowards enlargement of the database (Teel al. 2003), progress was anticipatedthat the databasewould et al. 1999).The allozymedatabase grewto zymedatabasewas slow for multi-jurisand multipledatabases allow for a wide varietyof management •ncludecomprehensive coverageof popu- dictionalfisheries, andresearch applications onall lifehistory lations rangingfrom California through proliferated. stages throughout the species rangein both Concerns regarding thesemultipleand Alaska with representative populations freshwater and marine environments. wereaddressed at fromRussia(Teel et al. 1999). Although independentdatabases Box 1. Definitionof termscommonlyusedin geneticstockidentification of Pacificsalmon. Allele Allelic ladder An alternative formof a givengeneor DNAsequence that differsfromotherallelesin DNAsequence or phenotype. A pooledanddilutedaliquotof multiplePCRproducts thatoriginates frommultipleindividuals of knowngenotype. With the choiceof appropriate individuals, a ladderwith all the "rungs"necessary for successful interlaboratory allele AIIozyme AIlelicformof a proteinenzymeencodedat a givenlocus.AIIozymes areusually distinguished byprotein indexingcan be created. Locus(loci,plural) Microsatellite Neighborjoiningtree PCR SNP 542 electrophoresis and histochemical stainingtechniques. Thesitethat a particular geneor DNAsequence occupies on a chromosome. DNAsequences containing short(2-5 basepairs)repeatsof nucleotides (e.g.GTGTGTGT). A bottom-upclustering methodusedfor the creationof phylogenetic treesthat isbasedon the distancebetween eachpairof populations or taxa. Thepolymerase chainreactionor PCRamplifies a singleor few copiesof a pieceof DNAacross several ordersof magnitude,generatingmillionsof copiesof the DNA. Singlenucleotide polymorphism; DNAsequence variation occurring whena singlenucleotide (A,T,C, or G)differs betweenmembers of a species or withinan individual betweenpairedchromosomes. Fisheries ß VOL 32 NO 11 ø NOVEMBER 2007 o WWW.F•SHERIES.ORG METHODS AND MATERIALS Figure1. a. Distribution of locianalyzed across laboratories at the beginning of thestudyMostmarkers wereassayed inonlya singlelaboratory. Onlythreelociwereusedinmorethanhalfof thelaboratories. b. Electropherograms fromthesameindividual assayed ontwodifferent instruments. Identical sizestandards Identificationof loci andsizing algorithm wereused, yetdifferences insizeareseenthatreflect differences inamplification chemistfids andelectrophoresis hardware, polymer andrunning conditions employed indifferent Thefigureillustrates a significant difference intheestimated sizeof thealldes,inaddition to a During the firstyearof the study, laboratories. different relative sizeoftherepeatunit(i.e.,4.14versus 4.25bp;adopted fromMoranet al.2006). each contributing laboratory analyzed an identical set of 500 individuals drawn frompopulations rangingfromCalifornia, USA, to Kamchatka, Russia,for its own microsatellitelocuspanel. A total of 60 loci were evaluated,and "sponsorship" documents were created for each locus including description,source,analysis conditions,and locusvariability.A sub- Downloaded by [Oregon State University] at 16:22 19 September 2011 set of 25 loci drawn from the combined set was chosen based on qualitative selectioncriteria includingthe reliability of the PCR (polymerasechain reaction; seeBox 1) amplification,and, to a lesserextent, allelic sizerangeand number of alleles.Locusdiversityacrossthe rangewasgenerallynot a consideration since too few individualswere assdyed to reliablycomputestatisticalmeasures. Each laboratorytested the 25 loci on a subset of 96 individuals Microsatellite marker (N=62) 70 75 drawn from the original 500 individuals. Scoresfor these 96 individuals were submitted to the coordinatinglaboratory(Northwest Fisheries Science Center, National Marine FisheriesService). From this set, collaborators selected and tested the 15 most reliable loci for inclusion in the standardized baseline. A curator labora- ^ tory was identified for each locuswith responsibilityfor receiving,compiling, and distributingallele information. A 175.ool decision was also made to link alleles to tissuesfrom a specificindividualsalmon that wouldresidein eachlaboratory.An additional I 00.001 Fragment size (bp) set of locus documentation was prepared designating recognized alleles, assigningallelic nomenclature, Columbia; the second set was collected Constructionof the Database and designatingindividualsfor every from southeast Alaska. Allelic scoresfrom recognizedallele at each locus. These eachlaboratorywereadjustedto the recBaselinepopulations werechosenthat master allele lists were termed "curator ognizednomenclature,submittedto the represent all previously identifiedgenetic documents." coordinatinglaboratory,and compared lineagesof Chinook salmonfrom the across seven laboratories. Two additional southernend of the speciesrangenorth laboratories that had not been involved Genotypingevaluation to southeastAlaskawith focuson major in locus selection or the first blind test productionareasand likely contributors To assess the ability of eachlaboratory participatedin the secondblind test.The to PacificSalmonTreatyfisheries (Figure to accuratelyresolveall 15 loci, 2 setsof alleleswere not sequenced, so the abso- 2). Six laboratories(Alaska Department 96 "blind" sampleswere analyzed(first lute lengthwasunknown.Therefore,the of Fish and Game [ADFG]; Department andsecondblindsamples). Thesesamples modal score (the most common allele of Fisheriesand Oceans Canada [DFO]; River Inter-Tribal Fish had neverbeen previouslygenotypedin scoredacrossthe participatinglaborato- Columbia any of the laboratories. In both cases,test ries) wasdefinedto be the "correct"score. Commission[CRITFC]; Oregon State fish were drawn from marine fisheries and Concordance of scoresacross laboratories University [OSU]; SouthwestFisheries assumedto be highly mixed with broad or percent accuracyfor each locus for ScienceCenter, National Marine Fisheries representation of stocks.The samplesfor eachlaboratorywasbasedon this modal Service [SWFSC]; and Washington the first test were collected from British score. DepartmentofFishandWildlife [WDFW]) Fisherles ß VOL32 NO 11 ß NOVEMBER 2007 ß WWW.FISHERIES.ORG 543 Figure2. Collectionsitesfor Chinooksalmonincludedin the DNAbaseline.Thesesitesrepresentpopul•ionspotentially contributing to mixtures harvestedin PacificSalmonTreatyfisheries.Sitesare designatedby regionalgroupsfrom Table3. 36b 36a ..... 35 39 34d 34c 34a North America 33a 37b, c 26b 27b 37a, e 37d 26••_27d r31b 26c4d ', '•'•27c 31a Downloaded by [Oregon State University] at 16:22 19 September 2011 30a •b . •a"'•3a• '24c 1• 29a 29e •ld ½4a 16b 20a•lc 10a,1 10b 10c, 11b 20b 9b 9c 1314c 18b 6•'•_.• 7b 5a•5bc 31a2d I KM 0 contributed 250 500 to the construction of the database. Evaluation of the database Definition of regional groups. The database wasevaluatedforitsabilityto correctlyallocateto regionalgroupsdefined basedon a combinationof geneticsimilarity, geographic features,and management applications.Population estimatesare combinedinto regionalgroupsto provide the desiredaccuracy andprecisionof stock composition estimates. The initial list of regionalgroupsfor compositional analyses wasenlargedfromthoseusedfor the allo54.4 1,000 zymebaseline(Teel et al. 1999). In order to visualizehow the presentmicrosatellite data might be concordantwith and/or improveuponthislist,wecalculated pair- mixtures of Chinook salmon harvested in treaty fisheries.These simulations assessedwhether the baseline of microsat- ellite allelefrequencies providedsufficient wise chord distances(Cavalli-Sforzaand informationto identifyregionalgroupsin Edwards 1967) between all collections hypotheticalmixtures. and then usedPHYLIP (Felsenstein2004) Simulations wereperformedusingthe to createa neighborjoining tree (Saitou StatisticalPackage forAnalyzingMixtures and Nei 1987; seeBox 1). On sucha tree, (SPAM version3.7; Debevecet al. 2000). geneticallysimilarpopulationswill clus- Mixture genotypes wererandomlygenerter together,facilitatingthe definitionof ated from the baselineallele frequencies regionalgroups. assumingHardy-Weinbergequilibrium. Mixture simulation. Simulations were The baselineallelefrequencies wereparato accountfor samconductedto evaluate the application metricallyresampled of theseregionalgroupsto geneticstock pling variability.Eachsimulatedmixture identification(compositional analyses) of (N = 400) wasentirelycomposed (100%) Fisheries ß VOL32 No 11 ß NOVEMBER2007 ß WWW.FISHERIES.ORG of the regionalgroupunderstudywLth the overall concordance for the laboratoequal contribution of all populations riesat theseloci wasover 99% (Table 2b). wLthin the regionalgroup.Then each s•mulatedmixture was analyzedagainst Baseline construction Database construction Our primarygoalof developing a reph- the completebaseline.Bootstrapmeans for regionalgroupsand 90% confidence Dataprovidedbythecollaborators were •ntervals were derived from 1,000 simu- combined to createa largedatabase suited lationsper group.Regionalgroupswith to GSI compositionalanalysesof fishermean correct estimates of at least 90% are ies managedunder the Pacific Salmon considered highlyidentifiable in potential Treaty.A total of 220 samplecollections Downloaded by [Oregon State University] at 16:22 19 September 2011 DISCUSSION cable set of microsatellite loci for Chinook salmonwassuccessfully completed.With an averageabove90% correctallocation to regionalgroups,the DNA baseline providesan unprecedented resourcefor improved information and management m•xturesfrom treaty fisheries.Regional and 16,394 individual fish representing of Chinooksalmonduringtheir oceanand groups with meancorrectestimates lower 110putativepopulations wereincludedin than 90% can still be considered idenfreshwater life stages.AdditionallaboraVersion1.1 of the baseline(Table3, Figure toriesemployingvariouschemistries and nfiable in mixtures,but sourcesof mis2, Appendix1). All individualfishwere hardware will be able to use the database allocation should be considered when assayed across13 loci; however,failures throughthe useof voucherspecimens or LnteFpreting the results(e.g., Seebet al. at particularloci sometimes resultedin byusingtheallelicladders (seeBox1) that 2000). incompletemultilocusgenotypes. A target arecurrentlyunderconstruction forall loci was set of 144 individuals per population RESULTS (e.g.,LaHoodet al. 2002).Allelicladders with at least120 genotypes per locus.In consistof a collectionof allelescovering some populations, fewer than 120individ- a rangeof knownsizes Locus selection and evaluation that canbeusedas of microsatellites ualswere available,so actual samplesizes an internal measurement to standardize werenecessarily belowthe target. the data.Theseladdershold the promtse The number of alleles varied markof simplifying standardization aslaboratoedly across loci,rangingfrom9 (Ors9)to rieswill not needto analyzevoucherspecthandlingerrors,suchasmisalignment of individuals, werethe largestsourceof dis- 74 (Omm1080;Table 1). Acrossall loci, mensfor everyallele. All geographicregionsand genenc agreement amonglaboratories. In addition 487 alleles were observed.A voucher set froma singleindividualfor each lineageslikely to contributeto fisheries to data-handling errors,2 of the 15 loci of tissues selected andtestedstoodoutasbeingpoorly of the alleleswereidentifiedandprovided of CTC interestare representedin the standardized across3 or more laboratories. to all collaborators.For somealleles,insuf- presentbaseline,and thesedata should for complexfisherymtxThese 2 loci were removed,and the final ficient tissuewas available for sharing be appropriate tures that include diverse populations between all labs, so a second individual groupof 13lociwasidentified forinclusion locations.Although the tn the baseline(Table 1). Beyondthese known to exhibit the same allele was sub- from widespread The first blind test revealed that data- sourcesof conflict, the resultsof the first stituted as the voucher. Additional alleles currentbaselineis broad,it is not compre- bhndcomparison indicatedthat technical are expectedas coverageof the baseline hensive.Effortsare currentlyunderwayto increaselocal coverage.Expandedbasestandardization hadbeenaccomplished for exTpands. line datawill likely improvethe accuracy greaterthan 90% of the observed alleles of allocationto the regionalgroups,and, at each locus.Furtherefore,much of the Simulations in at leastsomecases, arelikelyto provtde remainingvariabilityamonglaboratories a finer scale of estimation (e.g.,sub-bastns wasdue to allelesbeingobserved in the Forty-tworegionalgroups weredefined within major river systems). Fine-scale bhndsamples that hadnot beenobserved basedon geneticsimilarity,geographic geographic allocation of mixtures and in the original96 samples. features,and managementapplications. The results of the second blind test potential assignment of individualfish Simulationswere performedto examine indicated that technical standardization to population-of-origin mayalsoprovide the accuracy andprecision of thisbaseline importantbiological andlife-history inforhad been accomplished for more than simulatingmixturescomposed of indimationsuchasmigrationtimingandpath95% of the observedalleles (Table 2a). vidualsdrawn entirely from populations waysand age-related changesin habitat However,a numberof low-concordance of a singleregion.Almostall regionswere use.Expansion of the baselineto include valuesfor specificloci indicatederrorsin definedas highly identifiablewith boot- populations the data from two of the laboratories.One throughout Alaska,Canada, strap means of 1,000 simulated mixtures and Russia is underway. In additionto laboratory obtaineda concordance of only above 90% to the correct regional group improved management, the baselinewdl 44%forOgo4,whereas otherlociprovided werethe Deschutes be usedto betterunderstand the biologyof by the samelaboratoryshowed98% con- (Table3). Exceptions andprovideinformationabout cordanceor more.Anotherlaboratory had Fall and Upper Stikine River regions the species 0% concordance at five loci and 82% at which had 89.5% and 84.5% mean correct effective population size, evolutionary allocations, respectively. In addition,the anddemographic another locus. Data for the other seven history,andpopulation loci fromthis secondlaboratorywereclose lower bound of the 90% confidence inter- boundaries. The results of the two blind tests were wasabovea 90%threshto the overallaverage.Cursorychecksin val forall regions old, with the exception of the Deschutes encouraging, but alsosignalleda need respective laboratories revealeddata-handlingerrors thatexplained mostofthedis- Fall,UpperStikineRiver,TakuRiver,and for continuedvigilancein errorcheckLng anddatamanipulation. On the onehand, AlaskaStikineRiver groups. crepancies. After correction of theseerrors Southeast Fisheries • vol 32 NO 11 * NOVEMBER 2007 o WWk•/ FISHER[ES,ORG 545 Table 1. Microsatellite locistandardized for Chinooksalmon.Reference,curatoragency, and observednumberof allelesare givenfor baselineVersion1.1. Locus Reference Curator agency• Observednumber of alleles Downloaded by [Oregon State University] at 16:22 19 September 2011 Ogo2 Ogo4 0ki100 Olsenet al. 1998 Olsenet al. 1998 DFOunpublished 2 ADFG WDFW DFO 26 20 48 Ornrnl080 Rexroad et al. 2001 SWFSC 74 Ots2Olb Ots208b 0ts211 0ts212 0•213 Ots3M Grieget al. 2003 Grieget al. 2003 Grieget al. 2003 Grieget al. 2003 Grieget al. 2003 Griegand Banks1999 ADFG CRITFC ADFG OSU OSU WDFW 52 54 43 36 48 19 Ors9 Banks et al. 1999 DFO 9 OtsG474 Williamson et al. 2002 CRITFC 19 Ssa408 Cairneyet al. 2000 NWFSC 39 laboratones were able to achievea very high degreeof concordance throughthe standardization process andvouchersample comparisons. Furthermore, two new laboratories that hadnot beenpreviously involved in locusselection,baselineconstruction, or the first blind test were able Laboratory abbreviations: OSU,OregonStateUniversity; SWFSC,SouthwestFisheries ScienceCenter--NationalMarine Fisheries Service;DFO,Departmentof Fishenes and OceansCanada;NWFSC,NorthwestFisheries ScienceCenter-NationalMarineFisheries Service; CRITFC,ColumbiaRiverInter-Tribal FishCommission; ADFG,AlaskaDepartmentof FishandGame;WDFW,WasNngtonDepartment of FishandWildlife. Personal communication, K. Miller,Department of Fisheries andOceansCanada,Nanaimo,BritishColumbia, Canada. to achieveveryhighconcordance without anysignificant errors.On the otherhand, significant errorsin datahandling werestfil apparentin initialblindsubmissions from two of the original laboratories. These results, althoughnot uniqueto microsatellite data(e.g.,White andShaklee1991), suggest that ongoingqualitycontroland errorcheckingprocedures bothinternally within each laboratoryand amonglaboratories are warranted to ensure database integrity. Table2. Proportional genotyping accuracy by laboratory and locusfor the secondblindtestof 13 microsatellite loci.Averages across locus and laboratory aregiven.A. Results assubmitted.B. Results aftercorrection for errors(seetext). A. Locus Lab I Lab 2 Lab 3 Lab4 Lab 5 Lab 6 Lab7 Lab8 Lab9 Average Ogo2 Ogo4 0.987 0.439 1.000 1.000 1.000 1.000 1.000 0.995 0.993 1.000 0.000 0.000 1.000 0.995 0.993 0.994 1.000 0.990 0.886 0.824 OkilO0 Ornrnl080 Ots2Olb Ots208b 0ts211 0.978 1.000 0.984 0.994 1.000 1.000 1.000 1.000 1.000 1.000 1.000 0.993 0.955 1.000 1.000 0.995 0.995 0.994 0.970 0.994 0.985 0.970 0.985 1.000 1.000 1.000 0.995 0.994 0.994 0.888 0.995 0.995 0.991 0ts212 0.989 0.996 Ors213 0.987 Ots3M 1.000 1.000 1.000 1.000 0.995 1.000 1.000 1.000 1.000 1.000 1.000 1.000 0.994 1.000 1.000 1.000 1.000 1.000 0.989 0.995 0.994 1.000 1.000 0.982 1.000 0.985 0.000 1.000 1.000 1.000 0.884 1.000 1.000 0.988 0.994 1.000 0.000 1.000 1.000 0.995 0.886 Ors9 1.000 1.000 1.000 1.000 1.000 0.823 1.000 1.000 1.000 0.980 OtsG474 0.995 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 0.999 Ssa408 0.987 0.929 1.000 1.000 1.000 1.000 1.000 1.000 1.000 0.991 Average 0.949 0.995 0.997 0.999 0.998 0.597 0.998 0.991 0.998 0.947 0.000 0.993 1.000 B. Locus Lab I Lab 2 Lab 3 Lab 4 Lab 5 Lab 6 Lab 7 Lab 8 Lab 9 Average Ogo2 Ogo4 0.987 0.994 1.000 1.000 1.000 1.000 1.000 0.995 0.993 1.000 0.988 0.968 1.000 0.995 0.993 0.994 1.000 0.990 0.996 0.993 0ki100 0.978 1.000 0.984 0.994 Ornrnl080 Ots2Olb Ots208b 1.000 1.000 1.000 1.000 1.000 1.000 0.970 1.000 0.994 1.000 0.995 1.000 1.000 0.938 1.000 0.994 1.000 0.992 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 0.993 1.000 0.995 0.995 0.985 0.970 1.000 0.995 0.995 0.995 0ts211 1.000 1.000 0.994 1.000 0.993 0.955 0.994 0.985 0.994 0.991 0ts212 0.989 1.000 1.000 1.000 1.000 0.989 0.995 0.994 1.000 0.996 0ts213 0.987 1.000 0.982 1.000 0.985 0.994 1.000 1.000 1.000 0.994 Ots3M 1.000 1.000 0.988 0.994 1.000 0.949 1.000 1.000 0.995 0.992 Ors9 1.000 1.000 1.000 1.000 1.000 0.979 1.000 1.000 1.000 0.998 OtsG474 0.995 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 0.999 Ssa408 0.987 0.929 1.000 1.000 1.000 1.000 1.000 1.000 1.000 0.991 Average 0.992 0.995 0.997 0.999 0.998 0.981 0.998 0.991 0.998 0.994 .546 Fisheries o VOL32 NO 'i 'i "NOVEMBER2007 "WWW.FISHERIES.ORG Table 3. Mean correctallocationsfrom 100% simulationsfor 42 regionalgroupsof Chinooksalmonpopulations.Bootstrapmeansof 1,000 simulatedmixturesandupperand lower90% bootstrapconfidence intervals aregiven.Numberof populations per regionisalsogiven. Downloaded by [Oregon State University] at 16:22 19 September 2011 Region number Number of populations Region name Bootstrap mean interval Lower Upper 0.916 0.908 0.980 0.971 0.962 0.998 1 2 3 CentralValleyfall CentralValleyspring CentralValleywinter 4 4 1 4 5 California Coast Klamath River 2 0.985 0.975 3 0.984 0.970 0.995 0.995 6 7 8 9 10 North California/SouthOregonCoast RogueRiver Mid OregonCoast North OregonCoast LowerColumbiaspring 1 2 3 3 3 0.972 0.941 0.944 0.957 0.971 0.956 0.912 0.918 0.933 0.952 0.987 0.968 0.968 0.979 0.987 11 Lower Columbia fall 3 0.973 0.956 0.987 12 Willamette 2 0.983 0.968 0.994 13 Mid Columbia Tule fall 1 0.969 0.949 0.986 14 Mid andUpperColumbiaspring 5 0.965 0.945 0.985 15 Deschutes fall 1 0.895 0.859 0.931 16 UpperColumbiasummer/fall 3 0.963 0.937 0.985 17 Snake River fall 1 0.946 0.915 0.974 18 19 20 21 SnakeRiverspring/summer WashingtonCoast SouthPugetSound NorthPugetSound 5 3 2 4 0.966 0.951 0.988 0.971 0.946 0.928 0.977 0.955 0.984 0.971 0.997 0.985 22 Lower Fraser 2 0.984 0.972 0.994 23 24 25 LowerThompsonRiver SouthThompsonRiver NorthThompsonRiver 2 3 2 0.983 0.976 0.978 0.971 0.962 0.964 0.994 0.990 0.990 26 Mid Fraser River 4 0.980 0.966 0.991 27 UpperFraserRiver 4 0.969 0.949 0.986 28 East Vancouver Island 2 0.979 0.966 0.991 29 30 West Vancouver Island South BC Mainland 5 2 0.990 0.975 0.980 0.961 0.997 0.988 31 32 Central BC Coast Lower Skeena River 3 2 0.960 0.950 0.940 0.927 0.978 0.971 33 UpperSkeenaRiver 3 0.950 0.925 0.971 34 Nass River 4 0.939 0.913 0.963 35 UpperStikineRiver 1 0.845 0.793 0.893 36 Taku River 4 0.920 0.884 0.954 37 Southern Southeast Alaska 5 0.969 0.950 0.986 38 SoutheastAlaska,StikineRiver 1 0.917 0.884 0.947 39 KingSalmonRiver 1 0.987 0.977 0.995 40 41 42 Chilkat River Alsek River Situk River 2 1 1 0.988 0.980 0.977 0.977 0.967 0.963 0.996 0.992 0.990 River 0.945 0.936 0.990 90% confidence into thesepopulations wasusedin the allozyme singleallozymegroupwasseparated studyof GuthrieandWilmot (2004). fall, spring,andwintergroups. Our simulationresultssuggest that the The neighbor-joiningtree (Figure3) presentdatabaseprovidesgreaterresolu- suggested that somegroupsthat had not Growth of and accessto the baseline uon than wasprovidedby the allozyme been distinct from one another based on baseline.Only the DeschutesFall and the allozymebaselinemight be resolvMore detailed analysesof the power UpperStikineregionsfell belowourtarget ablewith the presentmicrosatellite data. of the databaseare ongoing.We expect of 90% accuracy. Additionalpopulations to betterdefinelimitations Overall,37 regionalgroups wereresolvable futureanalyses and tests are needed to evaluate whether of the present databaseand to identify withtheallozyme dataset(Teelet al. 1999) thesegroupscan be accurately identified where the addition of baseline collections in thismicrosator,alternatively, shouldbe combinedwith while42 wererecognized powervia the additionof adjacentregionalgroups.The similarity ellite datasetdespitethat fact that more and increased new markers are most needed. The addipopulations were included in the allozyme amongthe Taku and $tikine river transboundarypopulations wasalsoobserved baseline than the microsatellite baseline. tion of singlenucleotidepolymorphism w•th the allozymedatasetof Teel et al. Gains in resolutionwere most apparent (SNP) markers(seeBox 1) which areeasand may reflectselective (1999), and a singlecombinedgroupfor in the California Central Valley where a ily standardized Power of the database Fisheries o VOL 32 r•O 11 o r•OV[r•B[R 2007 o WWW.FISHERIES,ORG 547 F,gure3 Nelghbordolmng treebasedonpairwise chorddistances between collections of Chinook salmon (Appendix 1) Numbers andvertical hnesonrlght•ndlcatereglonalgroupsasdeslgnatedlnTable3 Regional groupswlthonlyaslnglepopulatlonareln bold FeatherRiverHatcheryFall Battle Creek Fall StanislausRiver Fall ButteCreekFall 1,2,3 FeatherRiverHatchery Spring • • I I ButteCreek Spring Deer Creek Spring Mill Creek Spring -r• t r• I •l ri- I /'--1 / • • -- / • L • • , LowerDeschutes R•ver SacramentoWinter 15 LyonsFerryHatchery Fall 11 Hanford Reach Creek Summer/Fall WellsDamHatchery Summer/Fall Methow River Summer/Fall SolDucRiverSpring • I I • I • r• I 1•9 16 SpringCreekHatcheryFall 13 Cowlitz Hatchery Spring • • •Cow•tzHatchery Fall I Lewis •iver I-all I Sandy River Fail r I ,u, 11 LewisHatcherySpring Kalama HatcherySpring '•L •-• [-- I McKenzieHatcherySpring • NorthSantiam Hatchery Spring Eel River Fall Russian River Fall Chetco River Fall RiverSpring Klamath River Fall I HatcheryFall [• Trinity Trinity Hatchery Spring 4-9 Downloaded by [Oregon State University] at 16:22 19 September 2011 Cole RiversHatcherySpring ApplegateCreekFall CoquilleRiver Fall Siuslaw River Fall Nehalem River Fall Alsea River Fall Siletz River Fall h Queers River FallRiverFall I 19 Quillayute/Bogachiel Marble-NVl Hatchery ConumaHatchery I__ Robertson Hatchery Sarita Hatchery NitinatHatchery 28,29 Quinsam Hatchery White River HatcherySpring • Big Qualicum Hatchery River Spring I21 I 20 Soos Creek Hatchery FallNorth Fork Nooksack Suiattle-SkagitRiver Spring Skagit River Summer NorthForkStilliguamishRiverSummer Birkenhead River Hatchery Spring I 22 Chilliwack River Hatchery Fall -- TucannonHa..tchery Spring/Summer Upper Yakima Hatchery Spring Wenatche River spring Carson HatcherySpring Warm SpringsHatcherySpdng Secesh River Spring/Summer John Day RiverSpring Minam RiverSpring/Summer Rapid River HatcherySpring/Summer -Imnaha River Spring/Summer Clearwater I I Li• • I L • • Louis River Spring SpiusRiverHatchery Spring ' Nicola HatcherySpring ChilkoRiverSummer • • River Summer 14, 18 Quesnel River Summer I• ' r• 1 NechakoRiverSummer StuartRiverSummer 23 - 27 TorpyRiverSummer r• SalmonRiverSummer r• -Swift River Summer • MorkillRiverSummer / LowerThompsonRiver Fall I Middle Shuswap Hatchery Summer LowerAdamsRiverHatcheryFall Klinaklini River I itimat HatcheryPorteau Cove Hatchery 30, 31 Atnarko Hatchery Wannock River Hatchery Ecstall River 32 Kincolith River Chickamin River 34 __ Clear Creek King Creek -- 37, 38 CrippleCreek --- Andrews Creek Little Tahltan River River 35 Kowatua Creek Tatsatua Creek 4um River bine River BulkleyRiver Sustut River 32 - 34 -- KwinageessRiver Owegee River 0.01 541l King Salmon River Klukshu River I SitukRiver BigBoulder Creek Tahini River 39- 42 Fisheries o VOL32 NO 11 o NOVEMBER 2007 o WWVV.FISHERIES.ORG variation(S•nithet al. 2005;S•nithet al. in press)is presently In MayandSeptember of 2007,thePSCconvened a workshop focusing onthecurrentandfuturecapabilities, limitations, anduseof underway. stockidentification methods in oceansahnon management. As partof thecollaboration, a web-based application hasbeen genetic groupof scientists attended representing thedisciplines of developed to allowthefisheries andresearch community to access A diverse genetics, administration, population dynamics, the genotypedata,curatordocuments, andsupporting metadata. fisherymanagement, theory.Attendees weredividedinto fourworkgroups: The PacificSahnonCommission provideddevelopmental fund- andsampling manage•nent, logistics, and•nodeling/sampling. Reviews of ingto theNorthwestFisheries ScienceCenter,NationalMarine genetics, areavailable at the PSCwebsite(http://psc.org/). The FisheriesService, Seattle, Washington(www.nwfsc.noaa.gov/ the workshop willbepublished whencompleted in 2008. research/divisions/cbd/standardization.cfm), and a test versionof finalrecommendations Asgenetic approaches to fisheries management become morecomthewebapplication isnowbeingevaluated. The permanent locaweanticipate thegrowthof thisandsimilartypesof datationandsupport forthedatabase arecurrentlyunderdiscussion by monplace, thatarepublicly available through webapplications. Ukimately, the collaborating agencies and the PacificSahnonCommission. bases willprovide a wealthof information notonlyforfishUntil the final location of the databaseis established,the data thesedatabases butalsoforresearchers froma varietyofdisciplines are availableat the ADFG website(www.genetics.cf. adfg.state. eriesapplications, investigating diverse aspects of sahnonid life history, population genetak.us/publish/data/PSCchinookver 1_1.pdf). ics,andevolutionary theory. CONCLUSIONS Downloaded by [Oregon State University] at 16:22 19 September 2011 ACKNOWLEDGEMENTS Herewedescribed a largecollaborative studyto developa shared, Thisstudy wasresult ofa collaboration among threeNational Marine web-accessible database of DNA markers to allowmultipleagencies Fisheries Servicelaboratories•Northwest Fisheries ScienceCenter,Auke to conduct geneticstockidentification studies of Chinooksahnon. BayLaboratory, andSouthwest Fisheries Science Center;Washington Replicate scorh•g of the microsatellite lociwassuccessfully accom- Department of FishandWildlife;University of Idaho/Coltunbia River Fisheries Conmfission/Idaho Department of FishandGame; plishedamongninegeographically-dispersed laboratories. However, Inter-tribal StateUniversity; Canadian Department ofFisheries andOceans; thestudyalsorevealed thatongoing coordination anderror-checkingOregon and the Alaska Department of Fish and Game. The U.S. Fish andWildlife willberequired to ensure theintegrity ofthedatabase asit expands. Service, Anchorage, Alaska,andIdahoDepartment of FishandGame, Growthofthedatabase ispresentl v underwav viatheaddition ofnew Eagle, Idaho,participated in thesecond blindsample. Primary funding for collections andadditional SNP markers. The database is nowbeh•g thestudy wasprovided bytheU.S.Department of Commerce, NOAA, usedto estimate compositional analyses of PSC andotherfisheries NMFS,funds appropriated tofundthetechnical workrequired fortheU.S. around the Pacific Ocean. section ofthePacific Sahnon Treatyto implement theChinook Salmon Y'our' ,,-:,'., .y uu.:an.;:,;,reThe View our latest catalog at www.flo.qta_q.,com, or email us at: salesC•flo.vta.q,COm or call to discuss yourcustomtaggingneeds:(8•0) 843-1172 ' Fisheries ß VOL32 •O 11 ß NOVEMBER 2007 ß WWW.FISHERIES.ORG $49 Agreementregarding abundance basedmanUmvers•ty ofWashington, SeattleAvadable htemarkers forrainbowtrout(Oncorhynchus agementPartialfundingfor the U S F•shand at http//evoluuon geneucs washington edu/ myktss) AmmalGenetics 32 317-319 WfidhfeService wasprovided bytheU.S.Yukon phyhp/geune.htmL Saitou, N., andM. Nei. 1987.Theneighbor-joinRiverResearch andManagement Fund(USRM- Greig,C. A., andM. A. Banks.1999.Fivemulingmethod: a newmethod forreconstrucung 08-05)andto theUniversity of Washington by tiplexed microsatellite lociforrapidresponse phylogenetic trees.MolecularBiologyand JointInstituteforthe Studyof theAtmosphere runidentification of Califomia's endangered Evolution 4:406-425. andOcean(JISAO)underNOAA Cooperative winter Chinook salmon. Animal Genetics Seeb,L. W., C. Habicht,W. D. Templin,K. E. AgreementNA17RJ1232,Contribution1444. 30(4):318-320. Tarbox,R. Z. Davis, L. K. Bmnnian,and Significant matching fundswereprovided byall Greig, C., D. P. Jacobson, and M. A. Banks. J. E. Seeb.2000.Geneticdiversity ofsockeye 2003. New tetranucleotide microsatellitesfor participating institutions. We thank the many members of the Chinook Technical Committee ofCookInlet,Alaska, anditsapphcafine-scale discrimination amongendangered salmon of the PacificSalmon Committee for their vision ofpopulations affected by Chinooksalmon(Oncorhynchus tshawytscha). tiontomanagement and support for the studyand five anonymous Molecular the ExxonValdez oil spill.Transactions of the Ecology Notes3:376-379. reviewers for their valuable comments on the Guthrie, C. M., III, and R. L. Wilmot. 2004. American Fisheries Society 129:1223-1249 Genetic structure of wild Chinook salmon Shaklee, manuscript. J. B., T. D. Beacham, L. Seeb,andB. populations ofsoutheast Alaskaandnorthern A. White. 1999.Managing fisheries using REFERENCES BritishColumbia. Environmental Biology of genetic data:casestudies fromfourspecies of Fishes69:81-93. Pacificsalmon.Fisheries Research 43:45-78 Banks, M. A., M. S. Blouin, B. A. Baldwin, Hankin, D. G., J. H. Clark, R. B. Deriso,J. Shaklee, J. B., and S. R. Phelps. 1990. Operation V. K. Rashbrook, H. A. Fitzgerald,S. M. C. Garza, G. S. Modshima,B. E. Riddell, of a large-scale, multiagency programfor C. Schwarz, andJ. B. Scott.2005.Rcportof genetic stock identification.Amencan the expertpanelon the futureof the coded Society Symposium 7:817-830. satellites in Chinooksalmon(Oncorhynchus wiretagrecovery program forPacificsalmon. Fisheries Smith C. T., A. Antonovich, W. D. Templin, C. tshawytscha). Journal ofHeredity90:281-288; Pacific Salmon Commission Technical Report errataJournal ofHeredity 90(3):U1-U1. M. Elfstrom,S. R. Narmn, andL. W. Seen 18, Vancouver,British Columbia.Available Banks, M. A., V. K. Rashbrook,M. J. In press. Impactsof markerclass biasrelauve at: www. psc.org/pubs/CWT/EPfinalrepor t. Calavetta,C. A. Dean,andD. Hedgecock. pdf. to locus-specific variabilityon populauon 2000. Analysis of microsatelliteDNA LaHood,E. S.,P.Moran,J. Olsen,W. S. Grant, inferences in Chinooksalmon; a companson resolves geneticstructureand diversityof and L. K. Park. 2002. Microsatellite allele of SNPsto STRsandallozymes. Transactions Chinook salmon(Oncorhynchus tshawyts- laddersin two speciesof Pacificsalmon: oftheAmerican Fisheries Society. cha)in Califomia's CentralValley.Canadian preparation and field-testresults.Molecular Smith,C. T., J. E. Seeb,P. Schwenke, andL Journal of Fisheries andAquaticSciences 57: Ecology Notes 2:187-190. W. Seeb. 2005. Use of the 5'-nucleasereac915-927. Blankenship,and D. Hedgecock.1999. Downloaded by [Oregon State University] at 16:22 19 September 2011 Isolation and inheritance of novel micro- Beacham, T. D., K. J. Supernault, M. Wetklo, B. Deagle,K. Labaree,J. R. Irvine, J. R. Candy,K. M. Miller, R. J. Nelson,andR. Mariani, S., W. E Hutchinson, E. M. C. Hatfield,D. E. Ruzzante,E. J. Simmonds, T. G. Dahlgren,C. Andre,J. Brigham,E. tionforSNPgenotyping in Chinooksalmon Transactions of the American Fishenes Society134:207-217. Torstensen, and G. R. Carvalho. 2005. E. Withler.2003.The geographic basisfor Teel, D. J., P. A. Crane, C. M. Guthrie III, North Sea herring populationstructure population structure in Fraser RiverChinook A. R. Marshall, D. M. VanDoornik,W. revealedby microsatellite analysis. Marine salmon(Oncorhynchus tshawytscha). Fishery D. Templin,N. V. Varnavskaya, and L. Ecology Progress Series 303:245-257. Bulletin 101:229-242. W. Seeb. 1999. Comprehensive allozyme Marshall,A. R., M. Miller, C. Busack,and S. Cairney,M., J. B. Taggart,andB. Hoyheim. R. Phelps.1991.Geneticstockidentification database discriminates Chinook salmon 2000. Atlantic salmon(SdmosalarL.) and around the Pacific Rim. (NPAFC docuofthree1990Washington oceanand cross-species amplification in othersalmoniris. analysis Strait of Juan de Fuca Chinook salmon fisherment 440). AlaskaDepartment of Fishand Molecular Ecology 9:2175-2178. ies,GSI Summary Report91-1,Washington Game, Division of CommercialFishenes, Cavalli-Sforza,L. L., andA. W. Edwards.1967. SalmonResearch Anchorage. Phylogenetic analysis. Modelsand estima- Departmentof Fisheries, andDevelopment, Olympia, Washington. Weitkamp,L., and K. Neely. 2002. Coho tionprocedures. American Journal ofHuman Mobrand,L E., J. Bart, L. Blankenship, D. Genetics 19:233-257. salmon(Oncorhynchus kisutch) oceanmigraE. Campton,T. T. P. Evelyn,T. A. Flagg, Cronin,M. A., W. J. Spearman, R. L. Wilmot, tionpatterns: insight frommarinecoded-wuce C. V. Mahnken,L. W. Seeb,P. tL Seidel, J. C. Patton, and J. W. Bickham. 1993. tagrecoveries. CanadianJournalof Fishenes Mitochondrial DNA variation in Chinook andW. W. Smoker.2005.Hatchery refonn 59:1100-1115. inWashington State:principles andemerging andAquaticSciences (Oncorhynchus tshawytscha) andchumsalmon White,B. A., andJ. B. Shaklee.1991.Needfor Fisheries 30(6):11-23. (O. keta) detectedby restrictionemyme issues. replicated electrophoretic analyses in mulMoran, P., D. J. Teel, E. S. LaHood, J. Drake, analysis of polymerase chainreaction(PCR) tiagency genetic stock identification (GSI) and S. Kalinowski.2006. Standardising products. CanadianJournalof Fisheries and examplesfrom a pink salmon multi-laboratory microsatellite datain Pacific programs: AquaticSciences 50:708-715. sahnon: an historical view of the future. (Oncorhynchus gorbuscha) GSI fisheries study CTC (Joint Chinook TechnicalCommittee). ofFreshwater Fish15:597-605. CanadianJournalof Fisheries andAquauc 2005. Annual exploitationrate analysis Ecology Nelson, tL J., M.P. Small, T. D. Beacham, Sciences48:1396-1407. (05)-3,Dec.28, 2005.Reportto the Pacific and K. J. Supernault.2001. PopulationWilliamson, K. S., J. F. Cordes,andB. May. structure of Fraser River Chinook salmon Salmon CommissionVancouver,British 2002. Characterization of microsatellite loc• tshawytscha): ananalysis using in Chinooksalmon(Oncorhynchus Columbia,Canada.Availableat: www. psc. (Oncorhynchus tshawytsDNA markers. Fishery Bulletin org/publications_tech_techcommitteere. microsatellite cha) and cross-species amplification in other 99:94-107. Debevec,E. M., IL B. Gates,M. Masuda,J. salmoniris. Molecular Ecology Notes 2:17-19 Pella,J. Reynolds,and L. W. Seeb.2000. Olsen,J. B., P. Bentzen,andJ. E. Seen 1998. Wirgin, I. I., andJ. R. Waldman.1994.What SPAM (Version3.2): Statisticsprogram Characterization of seven microsatellite loci 19(7):16-27 frompinksalmon. Molecular Ecology DNA candoforyou.Fisheries for analyzing mixtures. Journalof Heredity derived Wright, J. M., and P. Bentzen. 1994 91:509-511. 7:1083-1090. Microsatellites: geneticmarkersfor the Rexroad, C. E., III, R. L. Coleman,A.M. Felsenstein,J. 2004. PHYLIP (Phylogeny future.Reviews in FishBiology andFishenes Inference Package) version 3.6.Distributed by Martin, W. K. Hershberger, andJ. Killefer. theauthor,Department ofGenomeSciences, 2001. Thirty-fivepolymorphicmicrosatel- 4:384-388. and model calibration, TCCHINOOK 550 Fisheries ß vOL32 NO11 ß NOVE•IBER 2007 ß •/I,q/VW, FISHERIES.ORG Appendix1 Chinooksalmonpopulations analyzed inth•sstudyandIncluded in baseline Version1 1 Runt•me,hatchery (H)orw•ld(W)ongln,lifestage,collection data, andanalys•s laboratory areDven.Lowercasedes,gnat•ons w•th•na reDoncorrespond to locations on F•gure1. RegionnumberRegionname Population Runtime• OriginLifestage Collection date Analysislaboratory 2 1 CentralValleyfall BattleCreek(a) FeatherHatchery fall(b) Stanislaus River(c) TuolumneRiver(d) Fa Fa Fa Fa SWFSC SWFSC SWFSC SWFSC 2 CentralValleyspring 3 4 W H W W Adult Adult Adult Adult 2002, 2003 2003 2002 2002 CentralValleywinter ButteCreek(a) Sp DeerCreekspring(b) Sp FeatherHatchery spring(c) Sp MillCreekspring(d) Sp Sacramento Riverwinter Wi W W H W W/H Adult Adult Adult Adult Adult 2002, 2003 SWFSC 2002 SWFSC 2003 SWFSC 2002,2003 SWFSC 1992, 1993, 1994, 1995, 1997, SWFSC CaliforniaCoast EelRiver(a) Fa W Adult 2000, 2001 5 KlamathRiver RussianRiver(b) KlamathRiverfall (a) TrinityHatcheryfall (b) Fa Fa Fa W W H Juvenile Adult Adult 2001 2004 1992 6 North California/ Downloaded by [Oregon State University] at 16:22 19 September 2011 1998, 2001, 2003, 2004 SWFSC SWFSC SWFSC SWFSC TrinityHatchery spring(c) Sp H Adult 1992 SWFSC Chetco Fa W Adult 2004 OSU Applegate (a) ColeRivers Hatchery (b) Coquille(a) Fa Sp Fa W H W Adult Adult Adult 2004 2004 2000 OSU OSU OSU Siuslaw(b) Fa W Adult 2001 OSU Umpqua(c) Sp W Adult 2004 OSU AIsea(a) Nehalem(b) Fa Fa W W Adult Adult 2004 2000, 2002-1, 2002-2 Siletz(c) Fa W Adult 2000 OSU OSU OSU 2004 2004 2004 2004 CRITFC CRITFC CRITFC CRITFC 2003 2002, 2004 WDFW OSU SouthOregonCoast 7 RogueRiver 8 Mid OregonCoast 9 NorthOregonCoast 10 LowerColumbia spring 11 LowerColumbiafall 12 13 14 CowlitzHatchery spring(a)Sp KalamaHatchery spring(b)Sp LewisHatchery spring(c) Sp CowlitzHatchery fall (a) Fa H H H H Lewisfall (b) Sandy(c) Fa Fa W W Adult Adult Sp Sp Fa Sp Sp H H H H W Adult Adult UpperYakima(c) Sp WarmSprings Hatchery (d) Sp Wenatcheespring(e) $p H H W Adult, Mixed 1993, 1998, 2000 WDFW C RITFC WDFW LowerDeschutesRiver Fa W 1999-1, 1999-2, 2001,2002 CRITFC Su/Fa W 1999,2000-1,2000-2,2000-3,CRITFC W H 2001-1, 2001-2, 2001-3 1992, 1993, 1994 1993-1, 1993-2 WillametteRiver McKenzie (a) NorthSantiam(b) Mid Columbia Tulefall SpringCreek Mid and UpperColumbiaspringCarsonHatchery(a) JohnDay(b) 2002, 2004 2002, 2004-1,2004-2 2001,2002 2001,2004 Juvenile, Adult 2000-1,2000-2,2000-3, OSU OSU CRITFC CRITFC OSU 2000-4, 2000-5, 2000-6, 2004 15 Deschutesfall 16 UpperColumbia summer/fall HanfordReach(a) Methow Riversummer(b) Su/Fa Wells Dam (c) Su/Fa 17 18 SnakeRiverfall LyonsFerry SnakeRiverspring/summer ImnahaRiver(a) MinamRiver(b) RapidRiverHatchery (c) Sesech River(d) Tucannon (e) 19 Washington Coast Queets(a) Quillayute/Bogachiel (b) Fa Sp/Su Sp/Su Sp/Su Sp/Su Sp/Su Fa Fa W W W H W H W W 1998, 2003 2002, 2003 Adult Adult Adult Adult Adult CRITFC CRITFC 2002-1,2002-2, 2003-1, 2003-2 WDFVV 1998,2002,2003 CRITFC 1994,2002,2003 CRITFC 1997, 1999,2002 CRITFC 2001,2002,2003 2003-1,2003-2,2003-2 1996, 1997 1995-1, 1995-2, 1995-3, CRITFC WDFW WDFW WDFW 1996-1, 1996-2 20 SouthPugetSound SolDuc(c) SoosCreek(a) WhiteRiver(b) Sp Fa Sp Fisheries ß VOL32 NO 11 ß NOVEr•BER 2007 ß WWW.FISHERIES.ORG H H H Adult Adult adult 2003 1998-1, 1998-2,2004 1998-1,1998-2,2002 WDFW WDFW WDFW 55• 21 NorthPugetSound 22 LowerFraser NFNooksack (a) NFStilliguamish (b) Skagitsummer(c) Suiattle(Skagit)(d) Birkenhead River(a) Sp Su Su Sp Sp HNV HNV W W H adult adult adult adult Adult Chilliwack(b) Fa H Adult Nicola (a) Sp H SpiusRiver(b) Sp H LowerAdams(a) LowerThompson (b) MiddleShuswap (c) Clearwater(a) LouisRiver(b) Fa Fa $u Su Sp Chilko(a) Nechako(b) Quesnel(c) Stuart(d) MorkillRiver(a) SalmonRiver(Fraser)(b) Swift(c) TorpyRiver(d) BigQualicum(a) Quinsam(b) Conuma (a) Marbleat NVI(b) Nitinat(c) Robertson (d) Sarita(e) Klinaklini (a) PorteauCove(b) Atnarko(a) Kitimat(b) Wannock(c) Ecstall(a) Su Su Su Su Su Su Su Su 1999 1996 2001-1, 2001-2 1994 1995 1989 1998, 1999 1996 1997, 1999, 2001, 2002 Downloaded by [Oregon State University] at 16:22 19 September 2011 23 LowerThompson River 24 SouthThompson River 25 NorthThompson River 26 Mid FraserRiver 27 UpperFraserRiver 28 EastVancouver Island 29 WestVancouverIsland 30 SouthBCMainland 31 CentralBCCoast 32 Lower$keenaRiver 33 Upper$keenaRiver 34 NassRiver 35 36 UpperStikineRiver TakuRiver WDFW WDFW WDFW WDFW SWFSC 2003 1998 1999 DFO 1998 1999 OSU Adult 1996 1997, 1998 SWFSC H W H W W Adult Adult Adult Adult Adult 1996 2001 1997 1997 2001 DFO DFO DFO DFO DFO W W W W W W W W H H H H H H H W H H H H W Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult 1995, 1996, 1999, 2002 1996 1996 1996 2001 1997 1996 2001 1996 1996, 1998 1997, 1998 1996, 1999, 2000 1996 1996,2003 1997, 2001 1997 2003 1996 1997 1996 2000, 2001, 2002 DFO DFO DFO DFO DFO SWFSC DFO DFO DFO DFO DFO DFO DFO DFO DFO DFO DFO DFO DFO DFO DFO LowerKalum(b) W Adult 2001 DFO Babine(a) Bulkley (b) H W Adult Adult 1996 1999 DFO DFO Sustut(c) Damdochax(a) Kincolith(b) Kwinageese (c) Owegee(d) W W W W W Adult Adult Adult Adult Adult 2001 1996 1996 1996 1996 DFO DFO DFO DFO DFO LittleTahltanRiver KowatuaCreek(a) NakinaRiver(b) W W W Adult Adult Adult 1989 1990 1989 1990 1989 1990 OSU ADFG ADFG ADFG ADFG ADFG ADFG ADFG ADFG ADFG ADFG ADFG ADFG ADFG ADFG ADFG TatsatuaCreek(c) UpperNahlinRiver(d) Chickamin River(a) ClearCreek(b) CrippleCreek(c) Adult 1989 1990 W W W W Adult Adult Adult Adult 1989 1990,2004 1990 1993 1989 2003, 2004 1988 2003 37 SouthernSoutheast Alaska 38 39 Keta River(d) KingCreek(e) SoutheastAlaska,StikineRiver AndrewsCreek KingSalmonRiver KingSalmonRiver W W W W Adult Adult Adult Adult 1989 2003 2003 1989, 2004 1989, 1990, 1993 40 ChilkatRiver 41 42 AlsekRiver SitukRiver W W W W Adult Adult Adult Adult 1992, 1995,2004 1992, 2004 1989, 1990 1988, 1990, 1991, 1992 BigBoulderCreek(a) TahiniRiver(b) KlukshuRiver SitukRiver Runtimeabbreviations: spring(Sp),summer(Su),fall(Fa),andwinter(Wi) 2Laboratory abbreviations: OSU,OregonStateUniversity; SWFSC, Southwest Fisheries Science Center NationalMarineFisheries Service; DFO,Department of Fisheries andOceansCanada;CRITFC, Columbia River Inter-Tribal FishCommission; ADFG,AlaskaDepartment of Fish& Game;WDFW,Washington Depadmentof Fish& Wildlife. 552 Fisheries. vol 32 NO 11 . NOVEMBER 2007. WWW.FISHERIES.ORG