Wild Estonian and Russian sea trout (Salmo trutta) in Finnish coastal sea trout catches : results of genetic mixed-stock analysis

Wild Es onian and Russian sea ou ( Salmo u a ) in Finnish coas al sea ou ca ches: esul s o gene ic mixed-s ock analysis MARJA-LIISA KOLJONEN 1 , RIHO GROSS 2 and JARMO KOSKINIEMI 3 1 Finnish Game and Fishe ies Resea ch Ins i u e, Helsinki, Finland 2 Es onian Uni e si y o Li e Sciences, Ins i u e o Ve e ina y Medicine and Animal Sciences, Depa men o Aquacul u e, Ta u, Es onia 3 Uni e si y o Helsinki, Depa men o Ag icul u al Sciences, Finland Koljonen, M.-L., G oss, R. and Koskiniemi, J. 2014 . Wild Es onian and Russian sea ou ( Salmo u a ) in Finnish coas al sea ou ca ches: esul s o gene ic mixed-s ock analysis. – He edi as 151 : 177 – 195. Lund, Sweden. eISSN 1601-5223. Recei ed 1 Sep embe 2014. Accep ed 4 No embe 2014. Fo esponsible i she ies managemen o h ea ened species, i is essen ial o know he composi ion o ca ches and he ex en o which i she ies exploi weak wild popula ions. The h ea ened Es onian, Finnish and Russian sea ou popula ions in he Gul o Finland a e a ge s o mixed-s ock i she ies. The i sh may o igina e om i e s wi h a ying p oduc ion capaci ies, om di e en coun ies, and hey may also ha e ei he a wild o ha che y o igin. In o de o esol e he composi ion o Finnish coas al sea ou ca ches, we c ea ed a s anda dized baseline da ase o 15 DNA mic osa elli e loci o 59 sea ou popula ions a ound he Gul o Finland and es ed i s esolu ion o mixed-s ock analysis o 1372 cap u ed i sh. The baseline da ase p o ided su i cien esolu ion o eliable mix u e analysis a egional g oup le el, and also o mos o he indi idual i e s s ocks. The majo i y (76 – 80%) o he o al ca ch o igina ed om Finnish sea ou popula ions, 6 – 9% came om Russian and 12 – 15% om Es onian popula ions. Nea ly all Finnish ou in he ca ch we e o ha che y o igin, while he Russian and Es onian ou we e mos ly o wild o igin. The p opo - ion o i sh in he Finnish ca ches ha o igina ed om i e s wi h na u al p oduc ion was a leas one i h (22%, 19 – 23%). Two di e en spo ing pa e ns we e obse ed among he cap u ed ou , wi h a small and spa sely spo ed o m being ma kedly mo e common among indi iduals o Russian (28%) and Es onian o igin (22%) han among i sh assigned o a Finnish o igin (0.7%). Ma ja-Liisa Koljonen, Finnish Game and Fishe ies Resea ch Ins i u e, P.O. Box 2 FI-00791 Helsinki, Finland . E-mail: ma ja-liisa.koljo- [email p o ec ed] , ma ja-liisa.k[email p o ec ed] a e 1 Janua y 2015 He edi as 151: 177–195 (2014) © 2015 The Au ho s. This is an Open Access a icle. DOI: 10.1111/h d2.00070 The e a e cu en ly abou 101 i e s o b ooks d aining in o he Gul o Finland om Finland, Russia o Es onia in which he e is an anad omous ou popula ion ( Salmo u a L . ). O hese popula ions, 85 can be ega ded as na i e wild s ocks (ICES 2013). The emaining popula ions ha e been suppo ed by ha che y eleases. Fo abou one- hi d o he anad omous ou popula ions, he conse a ion s a us is e y poo , as o 29 popula ions he cu en smol p oduc ion le el is less han 5% o he po en ial smol p oduc ion le el o he i e . In addi ion, he conse a ion s a us is weak and unce ain o ano he 30 popula ions. In he Finnish Red Da a Book, he anad omous ou is lis ed as C i ically Endange ed, because na u al ep oduc- ion is uns able in mos Finnish Bal ic Sea popula ions due o in ensi e i shing ha also a ge s imma u e i sh, mig a ion obs uc ions and highly al e na ing wa e l ow le els in i e s ( KAUKORANTA e al. 2000, KALLIO-NYBERG e al. 2001, HEINIMAA e al. 2007, URHO e al. 2010). In o de o inc ease he escapemen o sea ou om he i sh- e y, he legal minimum ca ch size o sea ou was inc eased a he beginning o 2014 om 50 cm o 60 cm. In addi ion, in 2013, he legal minimum ca ch size o sea ou was inc eased in he Finnish go e nmen ally con- olled o sho e sea a ea o he Gul o Finland om 50 cm o 65 cm o adipose i n-clipped ou , all o he s a e comple ely p o ec ed om i she y and should be eleased i caugh . In Russia, a decla a ion o ou p ese a ion is in o ce so ha no legal ou i shing should occu . How- e e , despi e many o he i e s being si ua ed in he bo - de zone, poaching does occu . In he Es onian Red Lis o Th ea ened Species, he sea ou is lis ed as Nea Th ea - ened ( RED DATA BOOK OF ESTONIA 2008). In Finland, dam cons uc ion has been especially ac i e and se e al sea ou s ocks ha e been des oyed ( KALLIO- NYBERG e al. 2001). In o de o compensa e o he dec eased popula ion abundance and p oduc ion le els, a i i cial ep oduc ion in ha che ies and he elease o ea ed i sh o eggs in o i e s a e commonly p ac iced wi h he aim o e-es ablishing ex inc o enhancing weak popula ions. In addi ion, ha che y eleases o imp o e sea ou ca ches ha e also been widely used along he coas al a ea. In all, 293 000 sea ou smol s we e eleased in o he Gul o Finland in 2012. The majo i y o hese (74%, i.e. 216 000 smol s, mainly o Isojoki and Inga skilanjoki o i- gin) we e om Finnish eleases, 22% (64 000 smol s, only sea ou om he Ri e Luga) came om Russia and he 178 M.-L. Koljonen e al. He edi as 151 (2014) emaining 4% (13 000 smol s, mainly o Pudisoo o igin) we e om Es onian eleases (ICES 2013). This also e l ec s he app oxima e le els o ecen yea s. Es onia has announced i s aim o end sea ou eleases. The mixed-s ock i she ies in he Gul o Finland no only a ge i sh o i e s wi h a ying p oduc ion capaci- ies o sepa a e na ional o igins, bu also a mix u e o wild and ha che y i sh. This in many espec s esembles he si ua ion wi h A lan ic salmon in he Bal ic Sea ( KOLJONEN 2006, ICES 2013), al hough he ou i she y is mo e es ic ed o coas al a eas. Some mixing o anad omous ou om di e en coun- ies is known o occu , as agging expe imen s ha e gen- e ally shown abou 5 – 10% o he ou agged in Finland o be e u ned om he Es onian coas and some also om Russia. Co espondingly, sea ou agged in Es onia ha e o some ex en been ecap u ed in Finnish coas al wa e s. The coas al sea ou ca ch in 2012 was 13 300 kg in Es onia and 15 900 kg in Finland. In addi ion, Finland announced a o al ca ch o 3800 kg om i e s (ICES 2013). In Russia, wild sea ou popula ions a e ound in a leas 40 i e s o s eams. The majo i y a e si ua ed along he no heas e n coas o he Gul o Finland, bu he i - e s wi h he mos abundan smol p oduc ion a e in he sou he n Ri e Luga a ea (Fig. 1). The o al smol p oduc- ion o Russian i e s has been es ima ed o be a leas 10 000 – 15 000 smol s. Smol ap expe imen s indica e ha be ween 2000 and 8000 sea ou smol s o na u al o igin annually mig a e in o he sea om he Luga, he la ges Russian ou i e ( TITOV and SENDEK 2008). In Russia, sea ou eleases a e only ca ied ou in o he Ri e Luga and comp ise i sh o i s own i e sys em. In 2014, a o al o 48 500 one-summe -old and 43 500 wo- summe -old ju eniles we e eleased. In Es onia, sea ou popula ions a e ound in 45 i e s and b ooks in he Gul o Finland egion, o which 38 ha e wild popula ions (ICES 2013). Ri e s wi h highe 2 1 3 5 4 10 11 12 16 14 13 6 17 18 19 23 35 22 20 24 38-42 32 3334 47 48 49 50 51 52 53 54 56 57 58 59 55 43 44 45 46 Helsinki Tallin S . Pe e sbu g Gul o Finland 0 15 30 60 90 120 km FINLAND ESTONIA RUSSIA Bay o Vybo g Ka elian Is hmus Vybo g 36 37 7 89 25 26 27 28 29 30 31 21 A chipelago Sea Ca ch 1 Ca ch 2 Ca ch 3 Fig. 1 . The sampled b own ou i e s in Finland, Russia and Es onia. The colou o he i e indica es i s quali y as a spawning si e and po en ial en i onmen o b own ou . Red: i e is closed; blue: i egula ep oduc ion occu s; and g een: open i e wi h egula na u al p oduc ion o b own ou popula ions. The names and numbe ing o he i e s a e he ollowing: 1) Au ajoki, 2) Paimionjoki, 3) Pu ilanjoki, 4) Uskelanjoki, 5) Kiskonjoki, 6) Fiska sinjoki, 7) Inga skilanjoki, 8) Siun ionjoki, 9) Mankinjoki, 10) Espoonjoki, 11) Van aanjoki, 12) Sipoonjoki, 13) Koskenkyl ä njoki, 14) Kymijoki, 15) Isojoki (ha che y s ock, no in he map), 16) Summanjoki, 17) Vi ojoki, 18) U palanjoki, 19) San ajoki, 20) Vilajoki, 21) Te ajoki, 22) Rakkolanjoki, 23) Mus ajoki, 24) Kilpeenjoki, 25) R ö mp ö inpu o, 26) Myllyoja, 27) Koi is onpu o, 28) Pen il ä noja, 29) Kello-oja, 30) Lohijoki, 31) Papinoja, 32) Toi olanpu o, 33) No kopu o, 34) Jukkolanpu o, 35) Inojoki, 36) Pikku ammeljoki, 37) Vammeljoki, 38) Ty ise ä noja, 39) Hu inoja, 40) Te ijoki, 41) Huumosenoja, 42) Kuokkalanpu o, 43) Rajajoki, 44) Vo onka, 45) Sis a, 46) Ha lonka, 47) Luga, 48) P ü haj õ gi, 49) Kunda, 50) Toolse, 51) Selja, 52) Loobu, 53) Valgej õ gi, 54) Pudisoo, 55) Mus oja, 56) Pi i a, 57) V ä ä na, 58) Keila, 59) Vasalemma. The bo de s o Finnish adminis a i e ELY Cen es on he coas a e indica ed by black lines. App oxima e sea ou ca ch a eas a e shown as da k blue shaded a eas along he Finnish coas . The blue a ows indica e he p e alen wa e cu en s in he Gul o Finland. He edi as 151 (2014) Gene ic mixed s ock analysis o sea ou ca ches 179 da ase o analysing he o igin o Finnish sea ou ca ches in he coas al wa e s o he eas e n Gul o Finland. The i e s a e numbe ed om wes o eas clockwise ound he gul so as he i e numbe s a e showing he geog aphical p oximi y o he i e s (Fig. 1, Appendix 1 Table A1). Se en een o he i e sys ems we e en i ely on he Finnish side o he coas . Se en o he i e s c oss he Russian bo de , so ha he uppe eaches o he i e s a e loca ed in Finland and he lowe pa s d ain in o he sea in Russia. In addi ion, 23 sea ou popula ions om ela- i ely na i e i e s on he Russian coas and also om 12 i e s om he Es onian coas we e included in he baseline da a (Fig. 1). The Finnish sea ou popula ion samples we e ob ained om i e s discha ging in o ei he he nea by wes e n a ea, he A chipelago Sea o in o he Gul o Finland. In addi ion o he bo de i e s ( he no he n coas o he Bay o Vybo g), he Russian samples co - e ed he sou he n coas o he Bay o Vybo g, he Ka e- lian Is hmus, he no he n coas o he Bay o S . Pe e sbu g and pa o he sou he n coas o he Gul o Finland. Es onian i e s we e loca ed on he sou hwes - e n coas o he Gul . The ype and size o he i e s ma kedly a y among he s udied coun ies and a eas. In Finland, he wa e sheds a e ypically la ge and he i e s a e o en ou le s o la ge lake and i e sys ems. In Russia and Es onia, he i e s a e o en small and shallow, wi h he excep ion o he Ri e Luga sys em. The s a us o he i e en i onmen in all coun ies is classi i ed in o h ee ca ego ies, which a e shown in Fig. 1. Open i e s a e indica ed in g een, pa ly open in blue and i e s e ches abo e mig a ion obs acles a e shown in ed. P elimina y in o ma ion has also been used o classi y he popula ions acco ding o hei le el o o iginali y as o iginal (na i e), mixed h ough s ocking o in oduced, depending on hei s ocking his o y. The mos in e es ing and aluable popula ions om he man- agemen poin o iew a e hose ha a e anad omous, iable, o iginal and s ill gene ically di e se. A subs uc- u e analysis wi hin each Finnish i e sys em, in cases whe e se e al samples we e aken, was pe o med in K OLJONEN e al. (2013), and only anad omous popula ions we e included in he p esen s udy. All 12 Es onian base- line popula ions and mos , 16 ou o 23, Russian samples we e new o his s udy. The Finnish coas is di ided in o h ee go e nmen al adminis a i e sec o s acco ding o he Cen es o Eco- nomic De elopmen , T anspo and he En i onmen (ELY Cen es). These Cen es a e esponsible o he egional implemen a ion and de elopmen asks o he go e nmen . The i e s a e lis ed sepa a ely o each ELY Cen e (Appendix 1 Table A1; bo de s a e also shown in Fig. 1). smol p oduc ion a e si ua ed in he cen al pa o he no h Es onian coas (Fig. 1). Wa e shed-based analyses o he gene ic s uc u e o Finnish b own ou popula ions ha e also p e iously been conduc ed o na ional pu poses wi h bo h allozymes and DNA mic osa elli es ( KOLJONEN 1989, M ARTTINEN and KOLJONEN 1989, KOLJONEN e al. 1992, K OLJONEN and SAURA 1992, NUOTIO and KOSKINIEMI 1995, SAURA 2005, AALTONEN 2009, 2011). In addi ion, esea ch eams om o he Bal ic Sea coun ies ha e in es iga ed some es ic ed a eas o i e sys ems o he Bal ic Sea d ainage basin by using mic osa elli e ma ke s ( WAS and WENNE 2002, HANSEN e al. 2006, L EHTONEN e al. 2009, SAMUILOVIENE e al. 2009). How- e e , his s udy is he i s o co e he whole Gul o Finland coas . Fo esponsible i she ies managemen o h ea ened species, i is essen ial o know he composi ion o ca ches and he ex en o which i she ies exploi weak wild p o- duc ion. Mixed-s ock analysis based on gene ic da a has been used o decades in analysing he composi ion o Paci i c salmon ca ches in No h Ame ica ( SHAKLEE e al. 1999, BEACHAM e al. 2001, 2008) and he A lan ic salmon i she y in Wes G eenland ( REDDIN and FRIEDLAND 1999). I has also been used o e eal he composi ion o A lan ic salmon ca ches in he Bal ic Sea since 2000 ( KOLJONEN 2006, ICES 2013). A gene ic baseline da abase o mixed- s ock analysis has addi ionally been c ea ed o A lan ic salmon o he whole Eu opean ange ( GRIFFITHS e al. 2010). Fu he mo e, ca ch composi ion analysis has been u ilized o analyse s ock p opo ions in b own ou ca ches in he la ge, no he n, Finnish Lake Ina i ( SWATDIPONG e al. 2013). The aim o his s udy was o es he possibili ies o using mic osa elli e da a o mixed-s ock analysis o sea ou ca ches in he Gul o Finland. The goals we e o: 1) collec in e na ional baseline da a om all sea ou s ocks a ound he Gul o Finland; 2) es he esolu ion o his baseline da a se wi h simula ions o analyse s ock and s ock g oup p opo ions, and also he use ulness o indi- idual assignmen s o cap u ed i sh; 3) analyse he na ional con ibu ions o ou p oduc ion in he s udied Finnish ca ches; 4) analyse he p opo ions o wild and ha che y- p oduced i sh in he sea ou ca ch; and 5) compa e he esul s ob ained wi h Bayesian and maximum likelihood es ima ion me hods MATERIAL AND METHODS Fish sampling om i e s o c ea e he baseline da ase Al oge he , 4224 sea ou indi iduals om 59 wa e sheds we e sampled in he cu en s udy o c ea e he baseline 180 M.-L. Koljonen e al. He edi as 151 (2014) olume wi h 3 μ l o ex ac ed DNA, 5 μ l o ki mas e mix and p ime s wi h concen a ions and dyes as p esen ed in he Appendix 1 Table A2. PCR eac ions we e ca ied ou in PTC200 The mal Cycle s (MJ Resea ch), and he empe a u e p o i le o he PCR p og am was sugges ed in he Type-i Mic osa elli e ki manual. The annealing empe a u e was 56 ° C. The ampli i ca ion p oduc s we e sepa a ed by capilla y elec opho esis on AB3130 (in Finland) and AB3500 (in Es onia) Gene ic Analyze s (Applied Biosys ems, Fos e Ci y, CA), and he sizes o he mic osa elli e alleles we e de e mined using Genemappe e . 4.0 and e . 4.1 so - wa e, espec i ely (Applied Biosys ems, Fos e Ci y, CA), and manually checked. C oss-labo a o y calib a ion and s anda disa ion o allele sizes o all loci was conduc ed be ween labo a o ies om he Uni e si y o Helsinki, Depa men o Ag icul u al Sciences and he Es onian Uni e si y o Li e Sciences, Ins i u e o Ve e ina y Medi- cine and Animal Sciences, Depa men o Aquacul u e. This allowed he pooling o geno ype da a o es ablish a join baseline da ase , and enables he u u e analysis o all ca ch samples in bo h labo a o ies. S a is ical analysis Pai wise F s alues we e calcula ed wi h FSTAT e . 2.9.3.2. (Feb ua y 2002) ( GOUDET 1995, 2001) (  www2. unil.ch/popgen/so wa es/ s a .h m  ). Analysis o he di e ences be ween popula ions was based on geno ype equencies and was also es ed wi h FSTAT, which includes Bon e oni co ec ion o mul iple es s. Gene ic dis ances be ween sea ou popula ions we e calcula ed using Nei ’ s D A dis ances ( NEI e al. 1983). Phy- logene ic ees we e cons uc ed using a neighbou -joining (NJ) algo i hm ( SAITOU and NEI 1987, TAKEZAKI 1998) wi h Popula ions 1.2.32 so wa e ( LANGELLA 1999) (  h p:// bioin o ma ics.o g/ ∼ yphon/popula ions/  ). Boo s ap- ping wi h 1000 eplica es was used o es he s a is ical s eng h o he b anches. The gene ic dis ance ee was d awn wi h T eeView e . 1.6.6. ( PAGE 1996) (  h p:// axonomy.zoology.gla.ac.uk/ od/ ee iew.h ml  ). The ca ch composi ion was analysed wi h he Bayesian es ima ion me hod o mixed-s ock analysis using he BAYES p og am ( PELLA and MASUDA 2001). Gene ic in o ma ion on 59 sea ou popula ions a ound he Gul o Finland and A chipelago Sea om Finland, Russia and Es onia was a ailable as baseline popula ion da a. Ten i e a ion chains we e calcula ed o each ca ch es ima e. The p io o each po en ial baseline popula ion had a max- imum p opo ion o 71.0% in u n, and he o he s e enly 0.005%. The con e gence o he chains o he pos e io p obabili y dis ibu ion was es ed o each ca ch sample wi h con e gence s a is ics ( GELMAN and RUBIN 1992), included in he BAYES so wa e package. The las 1000 Sea ou sampling om i she y ca ches Fo he ca ch analysis, a o al o 1373 ou we e caugh by i she men, mos ly using bo om gill ne s (70%), bu also by od (17%) o yke ne (13%). Samples we e collec ed om bo h p o essional and ec ea ional i she y. The ca ch si es we e di ided in o h ee sec o s acco ding o he nea - es coas al municipali y (Table 1, Fig. 1). These ca ch sec- o s a e no he same as he ELY Cen e bo de s. All ca ch samples mos ly came om he eas e n pa o he Gul o Finland, be ween he ci y o Helsinki and he Russian bo - de , and ela i ely close o he Finnish coas . No ca ch samples we e a ailable om he wes e nmos pa o he coas om he a ea o he ELY Cen e o sou hwes Fin- land. Fishe y sampling was mo e e i cien in he eas e n- mos a ea (Table 1). The samples we e collec ed o e he yea s om 1996 o 2012 and hey do no hus ep esen any pa icula ime pe iod, bu a he p o ide an o e iew o he si ua ion in ecen yea s, when ha che y popula ion eleases ha e been qui e in ensi e in Finland. Mos sam- ples we e collec ed be ween yea s 2006 – 2012, and only samples less han 50 indi iduals in p e ious yea s. (Table 1). Fishe men also isually inspec ed i sh in hei ca ch and epo ed adipose i n-clipped i sh, and also i sh ha some- how de ia ed in hei appea ance. Some i sh we e epo ed o look mo e like salmon, being slende and ha ing ewe and smalle spo s han i she men we e used o ha e seen. These i sh we e iden i i ed om he indi idual assignmen da abase and also analysed sepa a ely. DNA me hods To al genomic DNA was ex ac ed om scale o issue samples in 95% alcohol using he DNeasy Blood & Tis- sue Ki (Qiagen) o acco ding o he simpli i ed me hod o L AIRD e al. (1991). Va ia ion was de e mined a 16 mic o- sa elli e loci (Appendix 1 Table A2). Howe e , locus SSa289 was omi ed om he s a is ical analysis, as i was no included in he Es onian baseline da a. Fo each sam- ple, wo mul iplex PCR eac ions we e pe o med using he Qiagen Type-i Mic osa elli e ki in a 10- μ l eac ion Table 1. Numbe s, sampling si es and main sampling yea s (  50 i sh yea 1 ) o he analysed Finnish sea ou ca ch samples. A ea Municipali y Loca ion n Main yea s 1 Vi olah i, Hamina, Ko ka Eas 656 2006 – 2012 2 Pyh ä ä , Ruo sinpyh ä ä , Lo iisa, Pe naja Middle 492 2009 – 2012 3 Po oo, Sipoo, Helsinki, Espoo, Ki kkonummi Wes 224 2010 – 2011 To al 1372 He edi as 151 (2014) Gene ic mixed s ock analysis o sea ou ca ches 181 10, wi h he co esponding mix u e sample sizes o 295, 413 and 590. RESULTS Gene ic di e en ia ion and dis ances be ween baseline popula ions The le el o di e en ia ion (as es ima ed by F s ) be ween pai s o baseline popula ions a ied om 0.002 o 0.437. Mos o he baseline popula ions we e signi i can ly di - e en ia ed om each o he , and s a is ically non-signi i - can di e ences in geno ype equencies (a e Bon e oni co ec ion o mul iple es s) we e only obse ed o wo Finnish popula ion pai s and one Es onian popula ion pai (F s alues o hese pai s a e p esen ed in ed in Table 2): Au ajoki – Fiska sinjoki (F s 0.009), Inga skilanjoki – Koskenkyl ä njoki (F s 0.013) and Pudisoo – P ü haj õ gi (F s 0.002). The geno ype equencies o hese popula ion pai s we e hus iden ical o each o he , and i sh ans e s o eleases a e known o ha e occu ed in all o hem. The di e ence was also only signi i can a he 5% nominal le el o wo e y simila popula ion pai s o i e s, Paimionjoki (2) – Siun ionjoki (8) in Finland and Hu - inoja (40) – Kuokkalanpu o (42) (Fig. 1) on he Ka elian Is hmus in Russia. The la e pai o i e s should ha e con ained na i e popula ions, bu a e geog aphically e y close o each o he (Fig. 1). All i e s numbe ed om 38 o 42 almos discha ge in o he same loca ion. In addi ion o he al eady men ioned e y simila popu- la ion pai s, F s alues smalle han 0.02 we e obse ed be ween hi een Es onian popula ion pai s, and also be ween occasional popula ion pai s in some o he geo- g aphically close i e s om o he a eas (all shown shaded in Table 2). All F s alues less han 0.02 we e, howe e , obse ed be ween popula ion pai s o he same geog aph- ical a ea. Sepa a ing hese e y simila popula ions on he basis o gene ic da a could be assumed o be di i cul , and hese popula ions a e unlikely o be dis inguished as indi- idual popula ions in mixed-s ock analysis, and we e hus o special in e es in he simula ion analysis. In mos o he cases wi h a e y high simila i y be ween popula ions, gene ic e ec s o ha che y eleases o known sou ces could be assumed acco ding o he his o y o ha che y i sh eleases. Among he Finnish popula ions, Au ajoki ou ha e been eleased in o he Ri e Fiska s- injoki and Koskenkyl ä njoki has been es ocked using Inga skilanjoki ou , which explains hei iden i y, while Isojoki ou ha e also egula ly been eleased in o he Ri e s Kymijoki and Summanjoki, which can be seen in he high simila i y be ween hem. In Es onia, Pudisoo and P ü haj õ gi ou ha e a leas also been mixed, in addi ion o some o he s, bu he geog aphical p oximi y he e is also high. In Russia, mixing o popula ions is no known o MCMC i e a ions o each 6000-i e a ion chain we e com- bined and used o desc ibe he pos e io p obabili y dis i- bu ions o he p opo ions o each baseline popula ion, and he eigh popula ion g oups ( o med based on he gene ic dis ances among baseline popula ions), as well as o he indi idual assignmen s. The Bayesian es ima ion was chosen o he es ima ion o he i nal esul s, as i has been shown in p e ious s udies o ou -pe o m he maxi- mum likelihood-based me hods ONCOR and SPAM, e.g. in es s whe e mixed samples o known o igin we e a ail- able ( GRIFFITHS e al. 2010, MORAN e al. 2014). Fo compa ison, he maximum likelihood es ima es o he same ca ches we e assessed wi h ONCOR so wa e (  www.mon ana.edu/kalinowski/So wa e/ONCOR. h m  ) ( KALINOWSKI e al. 2007, ANDERSON e al. 2008). The analysed sea ou baseline da ase o he Gul o Finland was e y la ge, including se e al simila se s o popula ions. I consis ed o nea ly 60 sea ou popula- ions, and hey o igina ed om a geog aphically ela i ely small a ea o some loca ions, which se s challenges o he es ima ion me hod. Fo example, when analysing a baseline da a se o 57 A lan ic salmon i e popula ions in he sou he n pa o he Eu opean ange, clea di i cul- ies we e me ega ding he es ima ion accu acy because o he gene ic simila i y o he popula ions, bu possibly also because o he small baseline sample sizes ( GRIFFITHS e al. 2010). Thus, he capaci y o he es ima ion me hod in gene al o sepa a e so many indi idual ou popula- ions and g oups was i s assessed by he simula ion op ions included in he ONCOR so wa e. The popula ion p opo ion es ima ion o indi idual i e popula ions was e alua ed wi h so-called 100% sim- ula ion, in which each baseline popula ion in u n con ib- u ed 100% o he analysed mix u e. The baseline samples we e he same as in he empi ical baseline. The mix u e sample size was se o 200 and 1300, which we e app ox- ima ely he smalles and la ges ca ch sizes used he e (224, 492, 656 and 1372, Table 1), and simula ions we e epea ed 100 imes o bo h indi idual popula ions and he eigh popula ion g oups used he e as epo ing g oups. The accu acy o indi idual assignmen was es ed wi h he lea e-one-ou p ocedu e and esul s we e ob ained simila ly o indi idual popula ions and epo ing g oups. In addi ion, he accu acy o he es ima ion was assessed by a simula ion op ion called ‘ Realis ic i she y simula- ion ’ , wi h simula ed mix u es o 200 indi iduals so ha each o he baseline popula ions in a se o 20 popula ions con ibu ed 5% o he simula ed ca ch. The esul s o hese simula ions a e combined in o one able. In addi ion, he op ion ‘ Th ee way e o decomposi ion: Fishe y, Loci, Baseline ’ , was used o analyse he sou ce o obse ed unce ain y o each baseline popula ion by se ing he numbe o indi iduals in each popula ion o be 5, 7 and 182 M.-L. Koljonen e al. He edi as 151 (2014) Table 2. Gene ic di e en ia ion o he mos simila baseline popula ions when measu ed as pai wise F s alues. Values less han 0.01 we e s a is ically non- signi i can and a e highligh ed in ed, and all obse ed alues less han 0.02 a e highligh ed in o ange. Only columns and ows wi h alues less han 0.02 a e shown . Pop 1 Au ajoki 7 Inga skilanjoki 14 Kymijoki 31 Papinoja 33 No kopu o 35 Inojoki 39 Hu inoja 44 Vo onka 48 Pühaj õ gi 49 Kunda 51 Selja 52 Loobu 56 Pi i a 57 V ä ä na 58 Keila 6 Fiska sinjoki 0.009 FIN 13 Koskenkyl ä njoki 0.150 0.013 RUS RUS 15 ISOJOKI 0.046 0.116 0.010 32 Toi olanpu o 0.107 0.171 0.138 0.020 35 Inojoki 0.068 0.152 0.101 0.039 0.015 37 Vammeljoki 0.086 0.153 0.099 0.061 0.032 0.017 39 Hu inoja 0.070 0.147 0.105 0.029 0.019 0.014 41 Huumosenoja 0.070 0.160 0.112 0.063 0.032 0.020 0.016 42 Kuokkalanpu o 0.094 0.165 0.119 0.029 0.027 0.024 0.012 45 Sis ajoki 0.065 0.125 0.090 0.074 0.041 0.042 0.046 0.017 46 Ha lonka 0.057 0.145 0.093 0.077 0.046 0.036 0.045 0.020 51 Selja 0.047 0.125 0.086 0.075 0.053 0.045 0.045 0.049 0.025 0.017 52 Loobu 0.071 0.142 0.107 0.082 0.057 0.058 0.064 0.067 0.034 0.011 0.020 EST 54 Pudisoo 0.056 0.153 0.097 0.085 0.067 0.058 0.061 0.071 0.002 0.032 0.028 0.033 56 Pi i a 0.064 0.128 0.096 0.086 0.055 0.052 0.051 0.054 0.041 0.032 0.019 0.036 57 V ä ä na 0.051 0.125 0.096 0.073 0.052 0.045 0.042 0.052 0.025 0.024 0.015 0.027 0.031 58 Keila 0.057 0.133 0.098 0.064 0.041 0.035 0.042 0.047 0.024 0.014 0.012 0.017 0.028 0.015 59 Vasalemma 0.045 0.124 0.081 0.058 0.043 0.034 0.036 0.040 0.023 0.013 0.011 0.017 0.018 0.014 0.010 He edi as 151 (2014) Gene ic mixed s ock analysis o sea ou ca ches 183 he ela i ely na i e Mankinjoki, Espoonjoki and Sipoon- joki popula ions om he nea by a ea could a leas pa ly be assumed o be a esul o hei common e olu iona y his o y. The ou wes e nmos i e popula ions o med he Uskelanjoki g oup, and hese i e s mos ly d ain in o he A chipelago Sea. The Russian popula ions g ouped e y p ecisely acco ding o hei geog aphical dis ances. The Bay o Vybo g popula ions o med a igh g oup, and wi hin i e en he subg oups om he no he n and sou he n coas o he bay could be sepa a ed, as wo popula ions, Myllyoja (numbe 25) and R ö mp ö inpu o (26) om he sou he n coas , o med a small subg oup o i s own (Fig. 2). All he Russian popula ions om he Ka elian Is hmus g ouped oge he , bu wi h some dis inc ion om popula- ha e occu ed, al hough some Ri e Luga ou ha e been eleased back in o he Luga i sel . When a phylogene ic ee was cons uc ed based on he gene ic dis ances be ween all popula ions, a clea and log- ical g ouping could be seen in he baseline da a, which ollowed su p isingly well he geog aphical dis ances be ween he popula ions and he o m o he coas line (Fig. 2), despi e he long his o y o eleasing ha che y popula ions in o he a ea. The excep ions o he geog aph- ical o de along he coas line came om he al eady known ha che y eleases o Isojoki and Au ajoki ha che y popula ions. In addi ion, he elease his o y explains he simila i y o he Inga skilanjoki ou wi h he Koskenky- l ä njoki and Van aanjoki popula ions, in o which i has been eleased. The simila i y o he Inga skilanjoki and 27Koi is on- pu o 29Kello-oja 28Pen ilänjoki 31Papinoja 32Toi olanpu o 30Lohijoki 33No kopu o 34Jukkolanpu o 38Ty ise änoja 40Te ijoki 39Hu inoja 41Huumosenoja 42Kuokka- lanpu o 35Inojoki 36Pikku- ammeljoki 37Vam- meljoki 19San ajoki 22Rakkolanjoki 21Te ajoki 23Mus ajoki 24Kilpeenjoki 20Vilajoki 17Vi ojoki (FIN) 25Römpö inpu o 26Myllyoja 18U palanjoki 43Rajajoki 44Vo onka 45Sis a 46Ha lonka 47Luga 49Kunda 52Loobu 50Toolse 51Selja 53Valgejõgi 57Vääna 58Keila 59Vasalemma 48Pühajõgi 54Pudisoo 55Mus oja 56Pi i a 7Inga - skilanjoki 13Koskenkylänjoki 9Mankinjokinjoki 11Van aanjoki 10Espoonjoki 12Sipoonjoki 14Kymijoki 15Isojoki 16Summanjoki 1Au ajoki 6Fiska sinjoki 5KiskoPe niönjoki 2Paimionjoki 8Siun ionjoki 3Pu ilanjoki 4Uskelanjoki Es onian s ocks Finnish s ocks Finnish-Russian bo de i e s Russia Ka elian Is hmus Russia Sou h G3 G8 G7 G5 G4 G2 G1 G6 18 33 46 99 10 100 55 50 15 5 94 709 48 24 10 99 6 40 10 63 57 27 7 85 31 20 24 72 19 11 9 11 22 16 30 18 19 42 1 67 52 18 36 13 59 42 75 30 23 13 9 13 43 5 44 3 Fig. 2 . Gene ic dis ances (Nei ’ s D A dis ance, neighbou -joining un oo ed ee) among Finnish, Russian and Es onian anad omous b own ou s ocks in he Gul o Finland. Wa e sheds a e numbe ed om wes o eas clockwise a ound he gul and wa e shed num- be s a e p esen ed oge he wi h he i e names. Boo s ap numbe s o he b anches a e shown as a pe cen age om 1000 epea s. 184 M.-L. Koljonen e al. He edi as 151 (2014) Es onian coas , a sligh subg ouping could addi ionally be seen, and popula ions om he Ri e s Keila and V ä ä na we e also e y simila . In all, i e majo g oups could be o med based on hei gene ic dis ances: Finnish popula ions (g oups 1 – 4), bo - de i e popula ions (g oup 5), eas e n Russian popula- ions (g oup 6), sou he n Russian popula ions (g oup 7) and Es onian popula ions (g oup 8) (Fig. 2, 3). De ia ing om his, he Finnish Ri e Vi ojoki, loca ed nex o he bo de i e s, g ouped wi h hem acco ding o i s gene ic simila i y. Consequen ly, he Bay o Vybo g g oup ac u- ally includes bo de i e s and he Ri e Vi ojoki om Finland nea he bo de . The Finnish popula ions, in u n, could be u he di ided in o ou g oups: A chipelago Sea a ea popula ions (1), Au ajoki ha che y ype popula- ions (2), Inga skilanjoki ype popula ions (3) and Isojoki ha che y ype popula ions (4). As mos o he cap u ed i sh o igina ed om Finnish popula ions, i ne -scale g ouping was necessa y in o de o gain mo e de ailed in o ma ion on he ca ch composi ion. These eigh popu- la ion g oups we e used as epo ing g oups in he ca ch analysis (Fig. 2), and his se ed as he i s ou pu le el o he o igin in o ma ion (Table 3). The loca ions o he i e s in he epo ing g oups a e indica ed in Fig. 3. Fou o he g oups (G1 o G4) a e ions no h and sou h o he Inojoki (35). The Inojoki pop- ula ion belonged o he sou he n g oup, which mainly includes i e s d aining in o he Bay o S . Pe e sbu g. The Russian Ri e s Papinoja (31) – Toi olanpu o (32) and No kopu o (33) – Jukkolanpu o (34) o med geo- g aphically close pai s. In addi ion, a e y igh g oup o i e small i e s (38 – 42) was also ound on he no he n coas o he Bay o S . Pe e sbu g, all o which we e di - i cul o dis inguish om each o he . The Ri e s Hu inoja (39) – Huumosenoja (41) and also Ty ise ä noja (38) – Te ijoki (40) o med close pai s, wi h Kuokkalanpu o (42) belonging o he same g oup wi h all o hem (Fig. 1, 2). The Ri e Rajajoki (Ri e Sies a joki) o med an in e - media y ype o he sou he n coas al Russian g oup, in which he Ri e Luga has he la ges smol p oduc ion. The ou i e s on he sou he n coas d ain in o wo small gul s, wo in o each, and hei gene ic simila i y also e l ec ed his pa e n. The Ri e s Vo onka (44) and Sis a (45) o med a pai , as well as he Ri e s Ha lonka (47) and Luga (48). In e es ingly, he Es onian ou popula- ions clea ly di e ed om his Luga ype o ou , and we e e y simila o each o he . Some e ec s o ha che y i sh eleases could be seen, as Pudisoo and P ü haj õ gi we e gene ically e y simila , and Pudisoo ou is known o ha e been eleased in o he P ü haj õ gi i e . Along he 2 1 3 5 4 10 11 12 16 14 13 6 17 18 19 23 35 22 20 24 38-42 32 33 34 47 48 49 50 51 52 53 54 56 57 58 59 55 43 44 45 46 Helsinki Tallin S . Pe e sbu g Gul o Finland 0 15 30 60 90 120km FINLAND ESTONIA RUSSIA Bay o Vybo g Ka elian Is hmus Vybo g 36 37 7 8 9 25 26 27 28 29 30 31 21 A chipelago Sea G1 G2 G3 G4 G5 G6 G7 G8 Fig. 3 . The loca ion o he eigh gene ic ca ch analysis epo ing g oups o he Finnish, Russian and Es onian sea ou s ocks in he Gul o Finland. The i e numbe ing is he same as in Fig 1. Adminis a i e ELY Cen e bo de s on he Finnish coas a e indica ed by black lines. He edi as 151 (2014) Gene ic mixed s ock analysis o sea ou ca ches 185 popula ion g oup p opo ion es ima es. When he accu- acy o indi idual assignmen was assessed wi h he lea e- one-ou p ocedu e, he likelihood o co ec assignmen was o e 90% o only 7 ou o he 59 baseline popula- ions. Fo 20 i e popula ions i was o e 80%, while o 26 popula ions he likelihood o co ec assignmen was s ill o e 75%. Fo ou Es onian popula ions (Keila, Kunda, V ä ä na and Pudisoo) i was less han 50%, again indica ing a e y high deg ee o simila i y among some o hem. In pa icula , he Pudisoo popula ion had a e y low likelihood o being co ec ly assigned, and i was mo e o en assigned o P ü haj õ gi han o i s own i e , which indica es ha i possibly does no e en dese e he s a us o a sepa a e popula ion. Howe e , when he likelihood o co ec assignmen o he indi iduals o hei own epo - ing g oup was es ima ed, on a e age up o 93% o he indi iduals we e assigned o he co ec popula ion g oup. Fo as many as 45 i e popula ions ou o 59 (76%), he pe cen age o co ec assignmen s was o e 90%. All indi- idual assignmen s a he g oup le el we e co ec , a leas a he 77% le el. The highes unce ain y occu ed be ween some Finnish ha che y popula ions. When simula ed mix u es wi h he same p opo ions o each popula ion we e analysed, a ia ion in es ima ion bias could be compa ed. Biases wi h a nominal alue o o e 1% o a 5% p opo ion occu ed o 16 ou o 59 baseline popula ions (Table 5). The unce ain y was high- es among he Es onian popula ions. In Finnish popula- ions, he p opo ion o Koskenkyl ä njoki ou was unde es ima ed in a ou o he Inga skilanjoki ou , om which i o igina es. On he Ka elian Is hmus, he sample om he i e Inojoki (baseline sample size n  98) popu- la ion seemed o ep esen well he ou o he a ea, as i s p opo ion was o e es ima ed a he cos o some o he comple ely on he Finnish coas , one g oup (G5) c osses he Finnish – Russian bo de , wo g oups a e loca ed in Russia (G6 and G7) and one g oup in Es onia (G8). Resolu ion o he baseline da ase When he so-called 100% simula ion o each baseline popula ion was conduc ed, he esul s showed ha a likeli- hood o co ec p opo ion es ima ion below 90% occu ed in he mos simila and ha che y ea ing-in l uenced Finn- ish popula ions (Au ajoki, Fiska sinjoki, Koskenkyl ä n- joki, Kymijoki), o which he s ocking his o y was known (Table 4). In addi ion, low alues we e also ob ained o ou na i e Russian popula ions on he Ka elian Is hmus, belonging o he igh popula ion g oup Vammeljoki (37), Ty ise ä noja (38), Hu inoja (39), Huumosenoja (41), Kuokkalanpu o (42), Vo onka (44, mixing wi h Sis a, 45) and Ha lonka (46, mixing wi h 47 Luga) on he Russian sou he n coas , and se e al Es onian popula ions (Valge- j õ gi, Pudisoo, Mus oja, Pi i a, V ä ä na, Keila and Vasa- lemma). The alue o he Pudisoo popula ion was especially poo , being only 0.09. Clea unce ain y is hus e iden in he indi idual popula ion es ima es o some o hese popula ions, and a leas he mos simila popula- ions should be g ouped in all po en ial g ouping op ions o ob ain eliable es ima es, especially o small ca ch sample analysis. Howe e , o all indi idual i e popula- ions, he likelihood o being co ec ly assigned o i s epo ing g oup was a leas 0.97 (Table 4.). I he mix u e sample size was inc eased o he maximum used o he o al ca ch o abou 1300 i sh, i did no essen ially a ec he esul . The e was conside ably mo e unce ain y in he indi- idual assignmen s o he i sh han o he popula ion o Table 3. Sea ou i e popula ions included in he epo ing g oups o he ca ch analysis. The mean F s alue o e s ock pai s wi hin each g oup is also p esen ed . G oup Ri e popula ions Coun y Mean F s 1 Au ajoki Au ajoki, Kiskonjoki, Fiska sinjoki FIN 0.079 2 Uskelanjoki Paimionjoki, Pu ilanjoki, Uskelanjoki, Siun ionjoki FIN 0.239 3 Inga skilanjoki Inga skilanjoki, Mankinjoki, Espoonjoki, Van aanjoki, Sipoonjoki, Koskenkyl ä njoki FIN 0.078 4 Isojoki Kymijoki, Isojoki, Summanjoki FIN 0.029 5 Bay o Vybo g Vi ojoki (FIN), U palanjoki, San ajoki, Vilajoki, Te ajoki, Rakkolanjoki, Mus ajoki, Kilpeenjoki, R ö mp ö inpu o, Myllyoja FIN/RUS 0.066 6 Ka elian Is hmus Koi is onpu o, Pen il ä njoki, Kello-oja, Lohijoki, Papinoja, Toi olanpu o, No kopu o, Jukkolanpu o, Inojoki, Pikku ammeljoki, Vammeljoki, Ty ise ä noja, Hu inoja, Te ijoki, Hu inoja, Kuokkalanpu o, Rajajoki RUS 0.055 7 Russia sou h coas Vo onka, Sis a, Ha lonka, Luga RUS 0.032 8 Es onia P ü haj õ gi, Pudisoo, Mus oja, Pi i a, Kunda, Toolse, Selja, Loobu, Valgej õ gi, V ä ä na, Keila, Vasalemma EST 0.028 192 M.-L. Koljonen e al. He edi as 151 (2014) Aal onen, J. 2011. Kiskonjoen – Pe ni ö njoen esis ö n s ä hk ö koekalas ukse uosina 2007 ja 2009 sek ä aimenkannan DNA-analyysi. – Salon seudun kalas usalue. Ande son, E. C., Waples, R. S. and Kalinowski, S. T. 2008. An imp o ed me hod o es ima ing he accu acy o gene ic s ock iden i i ca ion. – Can. J. Fish. Aqua . Sci. 65:1475 – 1486. Apa icio, E., Ga cia-Be hou, E., A aguas, R. M. e al. 2005. Body pigmen a ion pa e n o assess in og ession by ha che y s ocks in na i e Salmo u a om Medi e anean s eams. – J. Fish Biol. 67: 931 – 949. Beacham, T. D., Candy, J. R., Supe naul , K. J. e al. 2001. E alua ion and applica ion o mic osa elli e and majo his ocompa ibili y complex a ia ion o s ock iden i i ca ion. – T ans. Am. Fish. Soc. 130: 1116 – 1149. Beacham, T. D., Spils ed, B., Le, K. D. e al. 2008. Popula ion s uc u e and s ock iden i i ca ion o chum salmon Onco hynchus ke a om B i ish Columbia de e mined wi h mic osa elli e DNA a ia ion. – Can. J. Zool. 86: 1002 – 1014. Dege man, E., Leona dsson, K. and Lundq is , H. 2012. Coas al mig a ions, empo a y use o neighbou ing i e s, and g ow h o Sea ou ( Salmo u a ) om nine no he n Bal ic Sea i e s – ICES J. Ma . Sci. 69: 971 – 980. Gelman, A. and Rubin, D. B. 1992. In e ence om i e a i e simula ion using mul iple sequences. – S a . Sci. 7 : 457 – 511. Goude , J. 1995. FSTAT ( e . 1.2): a compu e p og am o calcula e F-s a is ics. – J. He ed. 86: 485 – 486. Goude , J. 2001. FSTAT, a p og am o es ima e and es gene di e si ies and i xa ion indices ( e . 2.9.3). G i i hs, A. M., Machado-Schia i no, G., Dillane, E. e al. 2010. Gene ic s ock iden i i ca ion o A lan ic salmon ( Salmo sala ) popula ions in he sou he n pa o he Eu opean ange. – BMC Gene . 2010: 11 – 31. Hansen, M. M., Bekke old, D., Jensen, L. F. e al. 2006. Gene ic es o a ion o a s ocked b own ou Salmo u a popula ion using mic osa elli e DNA analysis o his o ical and con empo a y samples. – J. Appl. Ecol. 43: 669 – 679. Heinimaa, P., Ju ila, E. and Paka inen, T. (eds.) 2007. I ä me en me i aimen y ö paja, (Bal ic Sea T ou Wo kshop). – Riis a- ja kala alouden u kimuslai os, Kala ja iis a apo eja n o 410. ICES 2013. Repo o he Bal ic Salmon and T ou Assessmen Wo king G oup 2013 (WGBAST), 3 – 12 Ap il, Tallinn, Es onia. – ICES CM 2013/ACOM:08. Kalinowski, S. T., Manlo e, K. R. and Tape , M. L. 2007. A compu e p og am o gene ic s ock iden i i ca ion. Manual. – Dep o Ecology, 310 Lewis Hall, Mon ana S a e Uni .  www.mon ana.edu/kalinowski/ONCOR/ONCOR_ Manual_21Oc 2007.pd  Kallio-Nybe g, I., Koljonen, M.-L. and Ju ila, E. 2001. Taimena las. – Kala u kimuksia 173. Kallio-Nybe g, I., Ju ila, E., Koljonen, M.-L. e al. 2010. Can he los mig a o y Salmo u a s ocks be compensa ed wi h esiden ou s ocks in he coas al i e s? – Fish. Res. 102: 69 – 79. Kauko an a, M., Koljonen, M.-L., Koskiniemi, J. e al. 2000. A las o Finnish i shes, English summa y. Dis ibu ion o lamp ey, b ook lamp ey, A lan ic salmon, b own ou , A c ic cha , whi e i sh, endace, g ayling, asp, imba, spined loach and bullhead – he dis ibu ion and s a us o s ocks. – Finnish Game and Fishe ies Res. Ins . Kocaba s¸ , M. and Ba s¸ ç ina , N. 2013. The e ec o salini y on spo ing ea u es o Salmo u a aban icus, S. u a a io such as Van aanjoki, Koskenkyl ä njoki and Inga skilan- joki i sel . The Isojoki ou o igina es om a i e ha does no d ain in o he Gul o Finland a all, bu u he o he no h, in o he Gul o Bo hnia. The use o his popula ion o ou eleases in he Gul o Finland could be econside ed in u u e, as i may well cause some gene l ow o he s ill na i e i e s. A ha che y b ood- s ock has al eady ounded om he bo de - i e , Mus a- joki, popula ion o po en ial use in eleases in he eas e n Gul o Finland ( PEUHKURI e al. 2014). The Rus- sian popula ions we e sligh ly mo e common in he eas e n ca ches. The baseline da ase collec ed in his s udy is highly aluable and will enable u u e ca ch analyses in all a eas o he Gul o Finland wi h easonable cos s. I hus o e s a use ul ool o i she ies managemen and conse a ion o he s ill na i e sea ou popula ions o he Gul o Finland a ea. Acknowledgemen s – This wo k was mainly i nanced by he INTERREG IV A P og amme p ojec HEALFISH ‘ Heal hy i sh s ocks – indica o s o success ul i e basin managemen ’ (2010 – 2013). A p opo ion o he Russian and Finnish baseline samples we e collec ed wi hin he INTERREG p ojec s ISKALT and ISKALT II ( SAULAMO e al. 2007, see sampling yea s in Table 1). Abou hal o he ca ch sample collec ion and analysis was i nanced by he Finnish – Russian p ojec ‘ Ri e s and i sh – ou common in e es ’ (RIFCI 2011 – 2014) unded by he Sou h- eas Finland – Russia ENPI CBC 2007 – 2013 P og amme. The ea lie analysed da a on Russian popula ions, collec ed du ing he ISKALT 2003 – 2007 p ojec s, was upda ed o 16 DNA mic- osa elli e loci om he p e iously analysed 10 loci da a se s, and his upda ing was i nanced by he RIFCI p ojec . Addi ional i nancing was p o ided by he Es onian Minis y o Educa ion and Resea ch (ins i u ional esea ch unding p ojec IUT8-2). We wa mly hank all ou colleagues om he Cen e o Eco- nomic De elopmen , T anspo and he En i onmen o Sou h- wes Finland, Uusimaa and Sou heas Finland, GosNIORKh, he Finnish Game and Fishe ies Resea ch Ins i u e and he Es onian Ma ine Ins i u e o he Uni e si y o Ta u o o ganizing and ealizing bo h i e popula ion and ca ch sampling. We a e also g a e ul o all he nume ous i she men who sampled hei ca ch i sh o he p ojec . F on ie gua ds made i possible o en e he on ie zone o sampling o he Finnish – Russian bo de i e s. A i Sau a and Aki Jana uinen p o ided aluable p ac ical in o - ma ion on he ou i e s and hei s a e in he Gul o Finland du ing he p ojec . Pekka V ä h ä n ä kki had a signi i can ole in highligh ing he impo ance o his p ojec o he bene i o common sea ou popula ions and i she men in all h ee coun- ies. We also wan o exp ess ou since e hanks o all he pa ies ha ha e p o ided unding o ou p ojec . D . Roy Siddall checked he English language. REFERENCES Aal onen, J. 2009. Elec o- i shing in he Uskelanjoki wa e cou se in yea 2006 and gene ic analysis o he b own ou ( Salmo u a L.) s ock. – Tu ku Uni . o Applied Sciences, Fishe ies and En i onmen al ca e, in Finnish. He edi as 151 (2014) Gene ic mixed s ock analysis o sea ou ca ches 193 Reddin, D. G. and F iedland, K. D. 1999. A his o y o iden i i ca ion o con inen o o igin o A lan ic salmon ( Salmo sala L.) a wes G eenland, 1969 – 1997. – Fish. Res. 43: 221 – 235. Sai ou, N. and Nei, M. 1987. 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Bayesian me hods o analysis o s ock mix u es om gene ic cha ac e s. – Fish. Bull. 99: 151 – 167. Peuhku i, N., Sau a, S., Koljonen, M.-L. e al. 2014. Cu en s a e and es o a ion o sea ou and A lan ic salmon popula ions in h ee i e sys ems in he eas e n Gul o Finland. – Wo king Pape s Finnish Game Fish. Ins . 26/2014. Red Da a Book o Es onia. 2008. – Commission o Na u e Conse a ion o he Es onian Academy o Sciences.  h p://elu ikkus.u .ee/ki jeldus.php?lang  eng & id  172944 & ank  70 & id_puu  172944 & ank_puu  70  accessed 27-08-2014. 194 M.-L. Koljonen e al. He edi as 151 (2014) Table A1. Analysed sea ou i e popula ion samples o he baseline da a om Finland, Russia and Es onia. The sampled i e , coun y o o igin, ibu a y o a ea, sampling yea , numbe o indi iduals and o iginali y o he sampled popula ion a e p esen ed. No Ri e Coun y T ibu a y Yea n O iginali y Wes coas , ELY Cen e o Sou hwes Finland 1 Au ajoki FIN 2006 37 In oduced 2 Paimionjoki FIN V ä h ä joki, Ka hunoja 2004, 2008 22 O iginal 3 Pu ilanjoki FIN 2011 15 O iginal 4 Uskelanjoki FIN Pi k ä koski, Kaukolankoski, Haukkalankoski, Hi olanjoki,Te il ä njoki 2007 57 O iginal, In oduced 5 Kiskonjoki FIN Pe ni ö njoki, Juo imenoja-Piilioja, Pakapy ö lin Lohioja 2008 50 O iginal Middle coas , ELY Cen e o Uusimaa 6 Fiska sinjoki FIN Main s eam 2010 50 In oduced 7 Inga skilanjoki FIN Main s eam, P ä hy elb ä cken, K ä ma s 2005 192 O iginal 8 Siun ionjoki FIN Passilankoski 2010 15 O iginal 9 Mankinjoki FIN Espoonka anonkoski, Gumb ö lenjoki 2005, 2008, 2010 133 O iginal 10 Espoonjoki FIN Main s eam, Glomsinjoki, Glimsinjoki 2008, 2010 72 O iginal 11 Van aanjoki FIN Van aankoski, Pi k ä koski, Ruu inkoski, Nuka inkoski, Longinoja 2010 207 Mixed, In oduced 12 Sipoonjoki FIN Ri ob ä cken, Byab ä cken 2010 46 O iginal 13 Koskenkyl ä njoki FIN Hamma o s, K a n o s, K ä kikoski, Sahakoski 2010 31 In oduced Eas coas , ELY Cen e o Sou heas Finland 14 Kymijoki FIN Main s eam 2006, 2010 96 In oduced 15 Isojoki FIN Lap ä ä in Isojoki, Ha che y s ock 2006 – 2008 98 Ha che y 16 Summanjoki FIN Main s eam 2003, 2008 19 Mixed 17 Vi ojoki FIN Saa asj ä enoja 2003, 2005, 2011 80 O iginal Bo de i e s 18 U palanjoki FIN/ RUS U palanjoenpu o (RUS), Main s eam (RUS, FIN) 1999, 2005, 2006, 2010, 2012 69 O iginal 19 San ajoki (Kal onjoki) FIN/ RUS Main s eam (RUS) 2005, 2006 50 O iginal 20 Vilajoki FIN/ RUS Main s eam (RUS) 2001, 2004, 2005, 2006 33 O iginal 21 Te ajoki FIN/ RUS Te ajoenpu o (RUS) 2004, 2005 48 O iginal 22 Rakkolanjoki FIN/ RUS Main s eam (RUS), Hanhijoki (RUS) 2004, 2005, 2006 109 O iginal 23 Mus ajoki FIN/ RUS Main s eam (FIN, RUS), Kananoja (RUS), Tupakkamyllynoja (FIN), Alhonpu o (FIN), P ö lkkyoja (FIN) 2004, 2005, 2006, 2007, 2008, 2012 438 O iginal 24 Kilpeenjoki FIN/ RUS Main s eam (RUS) 2001, 2006 16 O iginal Russian i e s 25 R ö mp ö inpu o RUS 2005 50 O iginal 26 Myllyoja RUS 2005 44 O iginal 27 Koi is onpu o RUS 2005 50 O iginal 28 Pen il ä noja RUS 2004, 2005 50 O iginal 29 Kello-oja RUS 2004, 2005 54 O iginal 30 Lohijoki RUS 2004, 2005 56 O iginal 31 Papinoja RUS 2004, 2005 46 O iginal 32 Toi olanpu o RUS 2004, 2005 52 O iginal 33 No kopu o RUS 2006 51 O iginal 34 Jukkolanpu o RUS Th ee close s eams 2004, 2005 148 O iginal Appendix 1 Con inued He edi as 151 (2014) Gene ic mixed s ock analysis o sea ou ca ches 195 No Ri e Coun y T ibu a y Yea n O iginali y 35 Inojoki RUS Main s eam and wo ibu a ies 2004, 2005 98 O iginal 36 Pikku ammeljoki RUS 2006 50 O iginal 37 Vammeljoki RUS 2004, 2005, 2006 50 O iginal 38 Ty ise ä noja RUS 2004, 2005 48 O iginal 39 Hu inoja RUS 2004, 2005 54 O iginal 40 Te ijoki RUS 2004, 2005 45 O iginal 41 Huumosenoja RUS 2004, 2005 50 O iginal 42 Kuokkalanpu o RUS Two close s eams 2006 23 O iginal 43 Rajajoki RUS Main s eam, Siesjoki 2006 21 O iginal 44 Vo onka RUS 2005 23 O iginal 45 Sis a RUS 2005 64 O iginal 46 Ha lonka RUS Ri e Tcho naja 2005 29 O iginal 47 Luga RUS Ri e Solka, R. Lemo zha, Vudoni s eam 2005 115 O iginal Es onian i e s 48 P ü haj õ gi EST 2010, 2011 176 Mixed 49 Kunda EST 2009, 2010 83 O iginal 50 Toolse EST 2011 49 O iginal 51 Selja EST 2010, 2011 116 Mixed 52 Loobu EST 2010, 2011 179 O iginal 53 Valgej õ gi EST 2010 39 Mixed 54 Pudisoo EST 2011 22 Mixed 55 Mus oja EST 2009 36 Mixed 56 Pi i a EST Lei a 2010 37 O iginal 57 V ä ä na EST 2011 68 O iginal 58 Keila EST 2010, 2011 66 O iginal 59 Vasalemma EST 2010, 2011 99 O iginal 4224 Table A2. Mic osa elli e loci used o sea ou analysis. Re e ences, mul iplexes, dyes and p ime concen a ions a e also indica ed. Locus Re e ence Mul iplex Dye P ime concen a ion 1 BS131 Es oup e al. 1998 MP 1 VIC 0.03 μ M 2 OneU9 Sch ibne e al. 1996 MP 2 VIC 0.03 μ M 3 SSa197 O’Reilly e al. 1996 MP 1 NED 0.02 μ M 4 SSa289 McConnell e al. 1995 MP 1 PET 0.30 μ M 5 Ssa407 Cai ney e al. 2000 MP 1 NED 0.15 μ M 6 SSa85 McConnell e al. 1995 MP 2 VIC 0.02 μ M 7 Ssosl311 Sle an e al. 1995 MP 2 NED 0.07 μ M 8 SSosl417 Sle an e al. 1995 MP 1 PET 0.04 μ M 9 SSosl438 Sle an e al. 1996 MP 2 VIC 0.07 μ M 10 SSsp1605 Pa e son e al. 2004 MP 2 NED 0.04 μ M 11 SSsp2201 Pa e son e al. 2004 MP 1 6-FAM 0.03 μ M 12 S 15INRA Es oup e al. 1993 MP 1 6-FAM 0.05 μ M 13 S 60lNRA Es oup e al. 1993 MP 2 PET 0.04 μ M 14 S 73lNRA Es oup e al. 1993 MP 1 VIC 0.04 μ M 15 S 85lNRA P esa and Guyoma d 1996 MP 2 6-FAM 0.40 μ M 16 S u 58 Po eaux 1995 MP 2 6-FAM 0.30 μ M Table A1. Con inued.