Selection on fish personality differs between a no-take marine reserve and fished areas

E olu iona y Applica ions. 2021;00:1–9. | 1wileyonlinelib a y.com/jou nal/e a 1 | INTRODUCTION Fishing- induced e olu ion and he consequences o popula ions ha e now been ex ensi ely documen ed (Kupa inen & Fes a- Bianche , 2017). Fo example, selec i e ishe ies may al e li e- his o y ai s in a popula ion by causing a shi owa ds ma u a ion a ea lie ages and smalle body sizes (Kupa inen e al., 2016; Olsen e al., 2004). Howe e , ishing- induced e olu ion o beha iou has ecei ed a less a en ion (Diaz Pauli & Sih, 2017). In e es ingly, g ow h a e can be ela ed o beha iou al exp ession, and a selec ion egime a ge ing la ge indi iduals may educe he o e all boldness in he popula ion compa ed wi h a selec ion egime whe e small indi iduals a e a ge ed (Bi o & Pos , 2008; Uusi- Heikkilä e al., 2015). Ha es ing may also selec di ec ly on beha iou (Uusi- Heikkilä e al., 2008). Fo exam- ple, passi e ishing gea can selec agains ai s such as s ong diel e ical mig a ion (Olsen e al., 2012) and la ge home anges (Alós e al., 2016) and lead o inc eased imidi y (A linghaus e al., 2017), while ac i e ishing gea such as awling may a ou bolde indi id- uals (Ande sen e al., 2018; Diaz Pauli e al., 2015). Mo eo e , since he ulne abili y o ce ain ha es condi ions may a y om one ish Recei ed: 31 Augus 2020 | Re ised: 24 Feb ua y 2021 | Accep ed: 23 Ma ch 2021 DOI: 10.1111/e a.13242 ORIGINAL ARTICLE Selec ion on ish pe sonali y di e s be ween a no- ake ma ine ese e and ished a eas Susanna Huneide Tho bjø nsen1,2  | E en Moland1,2 | Da id Villegas- Ríos3,4 | Ka inka Bleeke 1,2 | Hal o Knu sen1,2 | Esben Moland Olsen1,2 This is an open access a icle unde he e ms o he C ea i e Commons A ibu ion License, which pe mi s use, dis ibu ion and ep oduc ion in any medium, p o ided he o iginal wo k is p ope ly ci ed. © 2021 The Au ho s. E olu iona y Applica ions published by John Wiley & Sons L d 1Cen e o Coas al Resea ch, Depa men o Na u al Sciences, Uni e si y o Agde , K is iansand, No way 2Ins i u e o Ma ine Resea ch, Fløde igen, His, No way 3IMEDEA, Ins i u o Medi e áneo de Es udios A anzados (CSIC- UIB), Depa men o Ecology and Ma ine Resou ces, Ich hyology G oup, Espo les, Balea ic Islands, Spain 4IIM, Ins i u o de In es igaciones Ma inas (CSIC), Depa men o Ecology and Ma ine Resou ces, Fishe ies Ecology G oup, Vigo, Pon e ed a, Spain Co espondence Susanna Huneide Tho bjø nsen, Ins i u e o Ma ine Resea ch, Nye Fløde ig eien 20, N- 4817 His, No way. Email: susanna. ho bjo [email protected] Funding in o ma ion Resea ch Council o No way, G an /Awa d Numbe : 201917 PROMAR; FP7 ERA- Ne Biodi ERsA, G an /Awa d Numbe : 225592 BUFFER; Ma ie Sklodowska- Cu ie g an , G an /Awa d Numbe : 793627 (BEMAR); Eu opean Regional De elopmen Fund (In e eg IVa, Ma Gen II p ojec ); Coun y Go e no , Aus - Agde Abs ac Ma ine ese es can p o ec ish popula ions by inc easing abundance and body size, bu less is known abou he e ec o p o ec ion on ish beha iou . We looked o indi idual consis ency in mo emen beha iou s o sea ou in he ma ine habi a using acous ic eleme y o in es iga e whe he hey ep esen pe sonali y ai s and i so, do hey a ec su i al in ela ion o p o ec ion o e ed by a ma ine ese e. Home ange size had a epea abili y o 0.21, sugges ing ha i ep esen s a pe - sonali y ai , while mean swimming dep h, ac i i y and diu nal e ical mig a ion we e no epea able mo emen beha iou s. The e ec o home ange size on su i al di e ed depending on he p opo ion o ime ish spen in he ese e, whe e indi- iduals spending mo e ime in he ese e expe ienced a dec ease in su i al wi h la ge home anges while indi iduals spending li le ime in he ese e expe ienced an inc ease in su i al wi h la ge home anges. We sugges ha he di e si y o ish home ange sizes could be p ese ed by es ablishing ne wo ks o ma ine ese es en- compassing di e en habi a ypes, ensu ing bo h a he e ogenei y in en i onmen al condi ions and ishing p essu e. KEYWORDS acous ic eleme y, ha es selec ion, home ange, mo emen , pe sonali y, epea abili y, salmonids, spa ial ecology 2 | THORBJØRNSEN ET al. species o ano he (Killen e al., 2015), species- speci ic in o ma ion on beha iou al esponses o ishing es ic ions and p o ec ion will be im- po an o managemen . Pe sonali y is consis en indi idual di e ences in beha iou o e ime and h ough con ex s (Réale e al., 2007). Di e en beha iou al s a egies will be a ou ed in esponse o changes in a ange o en- i onmen al a iables, including ood a ailabili y, popula ion densi y and p eda o densi y, which is an impo an aspec in unde s anding hei main enance (Dingemanse & Réale, 2013). Assessing he he - i abili y o beha iou al ai s can be di icul , and epea abili y may be used as a p oxy (Doch e mann e al., 2015). P e iously, s udies on epea abili y o beha iou gene ally ha e been conduc ed in he lab- o a o y, bu mo e ecen ly esea che s ha e in es iga ed epea abil- i y o spa ial beha iou al ai s also in he wild (Ha ison e al., 2014; Villegas- Ríos e al., 2017). Such s udies a e impo an o unde - s anding how beha iou al a ia ion is main ained in na u e, which in u n may p o ide use ul inpu o adap ing conse a ion s a egies. Ma ine ese es ha e long been used as a conse a ion ool o p o ec agains deple ion om ishing (Les e e al., 2009). Howe e , i is unclea o wha ex en ma ine ese es may also help o p ese e beha iou al a ia ion wi hin popula ions by neu alizing ishing- induced selec ion (Baske & Ba ne , 2015). In e es ingly, ma ine ese es could d i e unan icipa ed selec ion on beha iou due o hei spa ial con igu a ion in ela ion o he spa ial mo emen s o he indi iduals, which migh ul ima ely e ode expec ed spillo e bene- i s (Villegas- Ríos e al., 2017). I is impo an hus o unde s and how selec ion may di e be ween ha es ed and p o ec ed a eas and o wha deg ee ma ine ese es may help in main aining he be- ha iou al di e si y wi hin popula ions, which ul ima ely ep esen esilience o en i onmen al change (Dingemanse e al., 2004). We used acous ic eleme y o quan i y mo emen beha iou and i s epea abili y o anad omous b own ou (Salmo u a) s ud- ied o up o 20 mon hs in ma ine habi a s in a Sou he n No wegian jo d. We hypo hesized ha sea ou mo emen beha iou , he e quan i ied as mon hly a e ages o he mo emen me ics home ange, mean swimming dep h, ac i i y and diu nal e ical mig a ion, was epea able among indi iduals and ep esen ed an aspec o hei pe sonali y. Ma ine ese es may al e he i ness o he indi iduals depending on how and whe e hey mo e. The e o e, we u he hy- po hesized ha selec ion on ou beha iou would di e be ween ished and p o ec ed a eas. 2 | MATERIALS AND METHODS 2.1 | S udy species The b own ou (Salmo u a) is a salmonid species in which ana- d omous popula ions a e called sea ou . Sea ou ha e a highly a iable li e his o y wi h some indi iduals spending only he sum- me a sea, while o he s spend mos o hei ime in ma ine a eas only e u ning o he i e o spawn du ing all (Kleme sen e al., 2003). Ma ine mig a ions a e mo i a ed by access o mo e ood, wi h impo an ade- o s being adjus men o di e en salini ies, inc eased ene ge ic cos o mo emen and a po en ially highe p e- da ion isk (Tho s ad e al., 2016). The balance o hese ade- o s is likely an impo an pa o he explana ion o he di e si y o mig a- ion s a egies wi hin popula ions (Tho s ad e al., 2016) and popula- ion di e en ia ion be ween s eams (Knu sen e al., 2001; Olsen e al., 2006). In No way, ishing o sea ou in ma ine habi a s is ec ea ional and pe mi ed all yea . Fishing can only be done using hook- and- line, excep o one mon h in summe whe e specialized aps a e pe mi ed in he sou he n pa o No way. The minimum legal size o sea ou in he ma ine habi a in No way is 35 cm. In he jo d, po en ial p eda o s o sea ou a e, among o he s, gulls, co mo an s (Phalac oco ax ca bo), ha bou seals (Phoca i ulina) and gadids, as epo ed om a s udy sys em in wes e n No way (Jonsson & Jonsson, 2009). 2.2 | S udy sys em and da a collec ion Mo emen da a we e collec ed in he T edes and jo d (3.8 km2, max dep h: 87 m) loca ed in sou he n No way be ween sp ing 2013 and all 2017 (Figu e 1). A eleme y a ay consis ing o 50 Vemco VR2- W ecei e s (VEMCO L d.) was deployed in he jo d, wi h he ecei e s being a ached o moo ings and kep a h ee me es dep h aided by subsu ace buoys ( o mo e de ails, see Villegas- Ríos, Réale e al., 2017). One ecei e was loca ed close o he spawning i e , Øs e åbekken, in o de o moni o i e mig a ions, and h ee ecei e s loca ed in he ou e pa o he jo d se ed as a ga e o de ec indi iduals dispe sing owa ds he ou e jo d and sea a eas. The high densi y o ecei e s ensu ed a good co e age o he jo d (see also Suppo ing In o ma ion, Tho bjø nsen e al., 2019). A ma ine p o ec ed a ea (1.5 km2) p ohibi - ing all ypes o ishing was es ablished wi hin he spa ial co e age o he eleme y a ay in 2012. Fishing is also p ohibi ed in Øs e åbekken and up o 100 m om he ou le o he s eam. Sea ou we e caugh a ound he cen e islands o he jo d in 2013 (Ap il: n = 3; May: n = 26; Sep embe : n = 24; No embe : n = 7), 2015 (June: n = 3; Oc obe : n = 14; No embe : n = 5) and 2016 (Ap il: n = 4; May: n = 7) using a beach seine, and also by elec o ishing in he spawning i e a 11 No embe 2016 (n = 23). Beach seine was cho- sen in an a emp o minimize sampling- induced selec ion o pa icula beha iou al ypes (Olsen e al., 2012). Elec o ishing was added as a complemen o inc ease sample size in 2016. Indi iduals we e anaes- he ized wi h clo e oil, and a ansmi e was inse ed in he abdomi- nal ca i y ( o de ails, see Olsen e al., 2012). We used Vemco V9P and V13P ansmi e s, which had a maximum ba e y li e o 508– 696 and 1292 days, espec i ely. Signals we e emi ed wi h a andom delay o 180 ± 70 s. Accu acy and esolu ion o dep h measu emen s we e ±2.5 m and 0.22 m, espec i ely, and max dep h was 50 m o mo e o he di e en ags. Sea ou we e no ex e nally ma ked. Fin clips we e aken o DNA analysis and p ese ed in 95% e hanol. In o al, 116 sea ou (mean body leng h: 337 mm, ange: 215– 635 mm) we e caugh , agged and moni o ed in he T edes and jo d du ing a 1669- day s udy pe iod (sp ing 2013– all 2017). A o al | 3 THORBJØRNSEN ET al. o 20 indi iduals we e excluded om he s udy due o ag mal unc- ion (n = 4), pos - su gical mo ali y (n = 5) o limi ed p esence in he s udy a ea (<14 days, n = 11). Time spen in he s udy anged om 1 o 20 mon hs. Ini ial da a explo a ion e ealed ha sex had no e ec on any beha iou al ai . 2.3 | Da a p epa a ion and es ima ion o beha iou al me ics De ec ions we e downloaded om he ecei e s and p ocessed using he VUE so wa e (VEMCO L d.), and u he da a p epa a ion and analyses we e done in he R en i onmen (R Co e Team, 2016). All de- ec ions a e p esumed dea h we e censo ed, which was de ined o ha e occu ed when con inuing de ec ions indica ed ha ho izon al and e ical mo emen had ceased (Olsen e al., 2012). No e ha his could also ep esen ansmi e loss. Fish we e de ined as dispe sed a e ha ing ollowed a di ec ional pa h ou o he ese e wi h inal de ec ions occu ing a he ou e mos ecei e s. Single de ec ions wi hin one day pe ish we e emo ed o elimina e po en ial code col- lisions and alse de ec ions, and abo e su ace dep h measu emen s we e de ined as NA. Fou ai s we e used o desc ibe he mo emen beha iou in he ma ine phase: home ange, mean swimming dep h, ac i i y and diu nal e ical mig a ion. Mon hly eplica es we e used o all ai s. Mon hly 95% home anges we e calcula ed using loca- ions based on posi ion a e ages (PAVs, cen es o ac i i y), ollow- ing Simp endo e e al. (2002). PAVs a e weigh ed a e age loca ions wi hin an a ay o ecei e s, based on he numbe o de ec ions a each ecei e du ing a speci ied ime pe iod (Simp endo e e al., 2002), in his case 30 min. Home anges we e hen calcula ed om PAVs using ke nel u iliza ion dis ibu ions (bandwid h = 60, ex en = 0.5) using he adehabi a HR package in R (Calenge, 2006). Dep h measu emen s we e a e aged o e mon hs a e emo ing eplica ed measu e- men s occu ing when a signal is de ec ed a mo e han one ecei e . Following F ei as e al. (2016), ac i i y was de ined as sho - e m changes in dep h and his was app oxima ed as he s anda d de ia- ion o dep h pe hou and hen a e aged o e mon hs. Diu nal e i- cal mig a ion was calcula ed as he di e ence in mean dep h om day o nigh wi hin a calenda day and hen a e aged o e mon hs. Day and nigh phases we e de ined by sola ele a ion da a ob ained om he Na ional Oceanic & A mosphe ic Adminis a ion (NOAA) h ough he map ools package in R (Bi and & Lewin- Koh, 2018). Only mon hs whe e he ish was p esen in he jo d o a minimum o 15 days (no necessa ily consecu i e) we e included in analyses. 2.4 | Repea abili y es ima ion Uni a ia e mixed- e ec s models we e i ed o each beha iou al ai using he nlme package (Pinhei o e al., 2018) in R. Fo mod- elling pu poses, home ange and ac i i y we e log- ans o med o mee no mali y assump ions o he esiduals. Mon hly a e ages o each beha iou al me ic se ed as eplica es o indi idual ish and indi idual sea ou iden i y was included as a andom e ec . We conside ed a ai o be epea able when he inclusion o he an- dom e ec signi ican ly imp o ed he model i . P o ided ha he andom e ec was suppo ed, epea abili y was calcula ed ollowing Dingemanse and Doch e mann (2013) as: whe e Vind0 is he among- indi idual a iance and V e 0 is he wi hin- indi idual a iance. Model selec ion was done in wo s eps: (1) selec ing he o e all model s uc u e by assessing i including (1) Repea abili y = V ind0 V ind0 +Ve 0 FIGURE 1 Map o he T edes and jo d (below) and i s loca ion along he No wegian Skage ak coas (abo e). The ma ine ese e in he cen e o he jo d is delinea ed wi h black lines. Blue do s ep esen ecei e loca ions, and ed do s ep esen cap u e loca ions 4 | THORBJØRNSEN ET al. he iden i y o he ish as a andom e ec and empo al au o- co ela ion be ween mon hs imp o ed he model (me hod = e- s ic ed maximum likelihood), ollowed by (2) selec ing he ixed e ec s s uc u e (me hod = maximum likelihood). Model selec ion was done using AIC- alues, and a minimum educ ion o >2 uni s was equi ed o assign signi ican imp o emen . When wo o mo e models ecei ed simila suppo , he model wi h he simples s uc u e was selec ed. Fixed e ec s included in he models we e body leng h (s anda dized o mean = 0, SD = 1), season (ca ego - ical a iable wi h ou le els, as de ined by he UK calenda wi h sp ing s a ing on Ma ch 1), sex and cap u e loca ion ( wo le els: jo d o i e ). Sex was de e mined using a sex- de e mining ma ke loci based on Eisb enne e al. (2014). 2.5 | Su i al A su i al cu e was gene a ed by compu ing a Kaplan– Meie es ima o o igh - censo ed da a (Cox & Oakes, 1984) using he ‘su i al’ package in R (The neau, 2015). Day o agging was se o 0 o all indi iduals. Fu he mo e, a Cox p opo ional haza ds e- g ession model was used o assess he ixed e ec s o home ange size and ese e use on su i al. Rese e use was included in mod- els ei he as he p opo ion o ime spen in he ese e gi en ha he ish was in he s udy a ea (calcula ed based on he loca ion o PAVs), o as cap u e loca ion ( wo le els: ese e o ished a ea), which se ed as a p oxy o co e a ea. Fish ha we e agged on he i e we e excluded om his analysis. Bo h home ange size and p opo ion o ime spen in he ese e we e calcula ed as he a e age o mon hly es ima es om agging un il dea h o he end o he s udy. Home ange sizes we e log- ans o med o mee no mali y assump ions. Addi ionally, he ixed e ec s ish leng h and season o cap u e ( wo le els: sp ing, as de ined by he sp ing agging season las ing om Ap il o June, and all, as de ined by he all agging season las ing om Sep embe o No embe ) we e also included in all models. Model selec ion based on AIC was done in wo s eps: (1) selec ing he bes model s uc u e ela ed o he main a iables o in e es (home ange size, p opo ion o ime spen in he ese e, agging loca ion), including in e ac ion e ec s be ween home ange size and p opo ion o ime spen in he ese e and home ange size and cap u e loca ion, and (2) selec ing he bes model ela ed o he addi ional co a ia es body leng h and season o cap u e. 3 | RESULTS Home ange size was a epea able mo emen ai ( epea abil- i y = 0.21, Table S1), while mean swimming dep h, ac i i y and diu - nal e ical mig a ion we e no (Tables S2– S4). Mean mon hly home ange size was 0.407 km2 ( ange: 0.065– 2.14 km2), inc eased wi h body leng h, and was la ge o ish caugh in he jo d han ish caugh in he i e (Table 1, Figu e S1). Home ange size was also a ec ed by season, being he la ges in sp ing, ollowed by all and summe , and he smalles in win e (Table 1). Analysis o mon hly mean swimming dep h (mean = 2.27 m, ange: 0.35– 9.44 m) showed ha ish caugh in he jo d swam deepe han ish caugh in he i e (Table 1, Figu e S1). Mean swimming dep h was also a ec ed by an in e ac ion be ween ish body leng h and season. Mean swimming dep h inc eased wi h body leng h and di e ed be ween seasons, wi h ish being loca ed a mo e shallow dep hs du ing all compa ed wi h all o he seasons. The in e ac ion be ween body leng h and season indica ed a s onge posi i e e ec o body leng h on mean swimming dep h in summe , ollowed by sp ing, win e and all (Table 1). Ac i i y (mean = 0.47 m, ange: 0.018– 3.67) inc eased wi h body leng h and was highe o ish caugh in he jo d (Table 1, Figu e S1). Ac i i y di e ed be ween he seasons, and ish we e mos ac i e du ing sp ing and summe , and leas ac i e du ing all and win e (Table 1). Diu nal e ical mig a ion, he di e ence in mean dep h om day o nigh (mean = 0.95 m, ange: −0.75 o 5.08), was la ge o ish caugh in he jo d han ish caugh in he i e (Figu e S1) and was a ec ed by an in e ac ion be ween body leng h and season (Table 1). Diu nal e ical mig a ion inc eased wi h body leng h and di e ed be ween seasons, wi h ish ha ing a la ge daily mo emen span du ing sp ing and summe han in win e and all. The in e ac- ion be ween body leng h and season indica ed a s onge posi i e e ec o body leng h on diu nal e ical mig a ion in sp ing and sum- me han in win e and all (Table 1). Including au oco ela ion led o signi ican imp o emen o all models wi h a beha iou al ai as he esponse a iable (Tables S1– S4). Es ima ed median su i al was 323 days (10.8 mon hs, Figu e 2). A his poin in he cu e, es ima ed su i al was 0.487 (95% CI 0.384– 0.617). The bes model p edic ing su i al included a e age mon hly home ange size, p opo ion o ime spen in he ese e and he in e ac ion be ween hese, in ad- di ion o season o agging (Table S5). The e ec o home ange size on su i al di e ed depending on he p opo ion o ime spen in he ese e (Figu e 3, Table 2). La ge home ange sizes inc eased su i al o indi iduals spending li le ime in he e- se e whe eas i dec eased su i al o indi iduals spending a la ge amoun o ime in he ese e. The e ec o home ange size on su i al wen om posi i e o nega i e when indi iduals spen mo e han 48% o hei ime inside he ese e. Fo ex- ample, model p edic ions showed ha i an indi idual spen 25% o i s ime in he ese e, an inc ease in home ange size om 0.265 km2 o 0.587 km2 would inc ease su i al a day 386 (las eco ded mo ali y e en ) by 20% (Figu e 3). In con as , model p edic ions showed ha i an indi idual spen 75% o i s ime in he ese e, an inc ease in home ange size om 0.265 km2 o 0.587 km2 would dec ease su i al a day 386 by 27%. A home ange size o 0.265 km2 co esponds o he 1s qua ile o home ange sizes, while a home ange size o 0.587 km2 co esponds o he 3 d qua ile o home ange sizes. Su i al was highe o ish agged in he all han ish agged in he sp ing. | 5 THORBJØRNSEN ET al. 4 | DISCUSSION Sea ou e ealed indi idual consis ency in home ange size o e a pe iod o se e al mon hs o e en yea s, e lec ing ha home ange can be conside ed an aspec o pe sonali y. Fu he , we ound ha home ange size a ecw ed su i al, and his ela ionship di e ed de- pending on he p opo ion o ime he ish spen inside he ese e. Fo indi iduals ha spen mo e han 48% o hei ime in he ese e, la ge home anges we e associa ed wi h dec eased su i al, while indi iduals ha spen less han 48% o hei ime in he ese e Response Pa ame e Es ima e SE d p- alue Home ange In e cep 12.1 0.224 272 <0.001 Leng h 0.134 0.0666 75 0.048 Cap u e loca ion, Fjo d 0.604 0.224 75 0.0086 Season, Win e −0.342 0.123 272 0.0058 Season, Sp ing 0.177 0.118 272 0.135 Season, Summe −0.145 0.111 272 0.193 Va iance, In e cep 0.3625 Va iance, Residual 0.6945 Mean dep h In e cep 0.678 0.331 324 0.0415 Leng h 0.127 0.152 324 0.404 Cap u e loca ion, Fjo d 0.996 0.326 324 0.0024 Season, Win e 0.487 0.219 324 0.0268 Season, Sp ing 0.881 0.203 324 <0.001 Season, Summe 0.827 0.190 324 <0.001 Season, Win e :Leng h 0.133 0.227 324 0.560 Season, Sp ing:Leng h 0.441 0.213 324 0.0395 Season, Summe :Leng h 0.671 0.206 324 0.0012 Ac i i y In e cep −1.67 0.174 322 <0.001 Leng h 0.251 0.0536 322 <0.001 Cap u e loca ion, Fjo d 0.445 0.175 322 0.0114 Season, Win e −0.0953 0.105 322 0.364 Season, Sp ing 0.317 0.0977 322 0.0013 Season, Summe 0.339 0.0896 322 <0.001 Diu nal e ical mig a ion In e cep 0.00313 0.231 279 0.989 Leng h 0.155 0.103 279 0.133 Cap u e loca ion, Fjo d 0.475 0.224 279 0.0348 Season, Win e 0.00867 0.156 279 0.956 Season, Sp ing 0.716 0.144 279 <0.001 Season, Summe 0.643 0.138 279 <0.001 Season, Win e :Leng h 0.0835 0.155 279 0.590 Season, Sp ing:Leng h 0.476 0.143 279 0.001 Season, Summe :Leng h 0.280 0.140 279 0.046 No e: Associa ed pa ame e es ima es, s anda d e o s (SE), deg ees o eedom (d ) and p- alues a e gi en. TABLE 1 Summa y o selec ed linea mixed- e ec s and lme models explaining mo emen beha iou in sea ou 6 | THORBJØRNSEN ET al. expe ienced inc eased su i al wi h inc easing home ange size. In o he wo ds, he i ness landscape o sea ou appea s o be in lu- enced by spa ial managemen , he e ep esen ed by a no- ake ma ine ese e. As discussed below, his sugges s ha ish beha iou migh e ol e in esponse o ishing and he e o e ce ain ishe y manage- men measu es. We ound ha home ange size had a epea abili y o 0.21, indi- ca ing ha 21% o he a ia ion in home ange size is a ia ion ha occu s among indi iduals. This is compa able o he beha iou al ai mean epea abili y o 0.37 o e all and 0.32 o ish p e iously e- po ed in a me a- analysis by Bell e al. (2009). Mo eo e , ou esul s con i m p e ious s udies showing epea able home ange in wild ish FIGURE 2 Righ - censo ed Kaplan– Meye su i al cu e o sea ou in he T edes and jo d. Red lines show median su i al a 323 days. Tagging day was se o ze o o all indi iduals. Ve ical ick ma ks indica e igh - censo ed e en s whe e an indi idual was no longe acked due o dispe sal o end o s udy o ba e y li e FIGURE 3 Su i al cu es o sea ou esul ing om he Cox p opo ional haza ds eg ession model wi h home ange size, p opo ion o ime spen in he ese e and season o agging as explana o y a iables. The ou panels show di e en combina ions o home ange size and p opo ion o ime spen in he ese e: (a) home ange size = 0.265 km2, p opo ion o ime in he ese e = 0.75; (b) home ange size = 0.587 km2, p opo ion o ime in he ese e = 0.75; (c) home ange size = 0.265 km2, p opo ion o ime in he ese e = 0.25; (d) home ange size = 0. 587 km2, p opo ion o ime in he ese e = 0.25. A home ange size o 0.265 km2 co esponds o he 1s qua ile o home ange sizes, while a home ange size o 0.587 km2 co esponds o he 3 d qua ile o home ange sizes. Season o agging had alue ‘ all’ in all su i al cu es 0.00 0.25 0.50 0.75 1.00 0100 200300 400500 600 Time Su i al p obabili y (a) Small HR, la ge p op. ese e 0.00 0.25 0.50 0.75 1.00 0100 200300 400500 600 Time Su i al p obabili y (c) Small HR, small p op. ese e 0.00 0.25 0.50 0.75 1.00 0100 200300 400500 600 Time Su i al p obabili y (b) La ge HR, la ge p op. ese e 0.00 0.25 0.50 0.75 1.00 0100 200300 400500 600 Time Su i al p obabili y (d) La ge HR, small p op. ese e | 7 THORBJØRNSEN ET al. (0.43 o A lan ic cod Gadus mo hua; Villegas- Ríos, Réale e al., 2017; 0.33 o bu bo Lo a lo a; Ha ison e al., 2014), sugges ing ha con- sis en mo emen beha iou may be a gene al pa e n o aqua ic o ganisms. Indi idual a ia ion in home ange sizes has also been shown in mammals, o example o moose (Alces alces; an Bees e al., 2011). Repea abili y was no de ec ed o any dep h- ela ed ai . No e ha ou mainly u ilized shallow dep hs, which means ha we may no ha e been able o de ec ine- scale di e ences in dep h use wi hin he gi en accu acy and esolu ion o dep h mea- su emen s. In u u e s udies o ine- scale dep h use, ags wi h highe esolu ion would be ad isable. Ou main inding is ha he e ec o home ange size on su i al di e ed depending on how much ime he ish spen in he ese e. Fo sea ou ha spen mo e han 48% o hei ime in he ese e su i al dec eased wi h inc easing home ange size. He e, a small home ange implies spending li le ime in he ished a ea (i.e. a isk o being ished). Consis en emo al o ish ha s and ou beyond ese e bounda ies may e en ually lead o selec ion agains la ge home anges in ma ine ese es (Villegas- Ríos, Moland e al., 2017). Fo sea ou ha spen less han 48% o hei ime in he ese e, su i al inc eased wi h inc easing home ange. No e ha his is he opposi e pa e n as p e iously ound by bo h Alós e al. (2016) who epo ed selec ion agains la ge home anges in ha es ed a eas, and Hä könen e al. (2014) inding ha being explo a i e is linked o an inc eased ulne abili y o angling in b own ou . O e all, he e is con as ing e idence on he ela ionship be ween home ange and su i ing he ishe y. Monk and A linghaus (2017) epo no e ec o swimming dis ance o ac i i y space on ulne abili y o cap u e by angling, while Olsen e al. (2012) epo inc eased ishe y su i al o ish ha displayed ex ensi e ho izon al shi s. In e es ingly, indi iduals ha exp ess mo e isky beha iou s in he wild, including explo a ion, expe ience inc eased su i al (Moi on e al., 2020). Risky beha iou s may lead o acqui ing mo e esou ces ollowed by an inc ease in na u- al su i al (Moi on e al., 2020). Ha ing a la ge home ange size may be one such isky beha iou . Fu he mo e, i is less ob ious why a la ge home ange mainly loca ed ou side he ese e yields be e su i al han a la ge home ange mainly loca ed inside he ese e. A s udy wi h eplica ed p o ec ed- unp o ec ed a ea pai s could help o in es iga e whe he he pa e ns ound in his s udy a e gene al pa e ns. These indings, combined wi h he ac ha home ange is epea able and he e o e likely pa ly he i able (Doch e mann e al., 2015), may en ail e olu iona y consequences o popula ions ha a e pa ially p o ec ed by ma ine ese es. Tha said, any local e olu iona y change will also depend on he le el o gene low (Leno mand, 2002). In acco dance wi h es ima es o how much addi i e gene ic a ia ion con ibu es o pe sonali y, Doch e mann e al. (2015) es ima ed ha he a io o he i abili y o epea abili y collec ed om li e a u e a e aged a 0.52 and anged om 0 o 0.96 (Doch e mann e al., 2015). Few s udies in es iga e he i abili y o beha iou in sea ou , bu in a labo a o y s udy on ad lu ial b own ou , Ko e e al. (2014) ound he i abil- i y o 0.14 (± 0.096) o he s ess esponse ‘ endency o eeze’, bu no he i abili y o boldness, explo a ion and agg ession. To he bes o ou knowledge, he e a e no es ima es o he i abili y o sea ou beha iou in he wild. Ou pape is he i s o p esen es ima es o epea abili y o sea ou beha iou in he wild, indica i e o addi i e gene ic a ia ion (Doch e mann e al., 2015). In addi ion o addi i e gene ic a ia ion, epea abili y may also e lec lea ning (Ad iaenssens & Johnsson, 2011) and indi idual a ia ion in u ilizing he e ogeneous en i onmen s (Bell e al., 2009). Hence, epea abili y o home ange size may also, o some deg ee, e lec indi idual di e ences in habi a use (Bell e al., 2009). Body leng h a ec ed all mo emen ai s, wi h la ge ish ha - ing la ge home anges, u ilizing a la ge ange o dep hs and ha - ing a highe ac i i y. As su i al was a ec ed by home ange size, his may imply co ela ed selec ion on body leng h. Howe e , body leng h did no a ec su i al di ec ly. Home anges we e he la ges in sp ing, and ish we e mo e ac i e du ing sp ing and summe han all and win e . This is in acco dance wi h sea ou in ensi ying hei ood sea ch as empe a u es inc ease du ing sp ing and summe (Kleme sen e al., 2003). Fish also swam deepe du ing sp ing and summe , which can be associa ed bo h wi h di e en habi a use and ha he ou seek ou colde wa e empe a u es op imal o g ow h when su ace empe a u es ise (Eldøy e al., 2017; K is ensen e al., 2018). Fish agged in he sea had la ge home anges, u ilized a la ge ange o dep hs and had highe ac i i y han ish agged in he i e . Median su i al in he wild was close o 11 mon hs, and su - i al was highe o ish agged in he all. The la e could be ex- plained by he upcoming spawning ascen , whe e sea ou will ecei e p o ec ion om ishing and expe ience a lowe p eda ion isk in he i e (Tho s ad e al., 2016). In gene al, ou su i al is highe in eshwa e as compa ed o sea (Solomon, 2006), and he du a ion o mig a ion a ies bo h wi hin popula ions and among popula ions and la i udes (Kleme sen e al., 2003). This implies ha yea ly su i al will a y subs an ially be ween i e sys ems. Re u n a es om 193 sea ou agged in he nea by i e S o el a (<5 km om ou s udy sys em) e ealed 40% su i al o ou spending one o wo yea s a sea (Ha alds ad, 2015). Su i al migh ha e been unde es ima ed due o ag exc e ion, which would ha e led indi iduals o be alsely de ined as dead. Also, he e migh be a nega i e e ec o agging on su i al. A s udy on gas ically agged salmonids ound ha small (9 mm) and la ge TABLE 2 Reg ession coe icien s, haza d a ios, s anda d e o (SE) o he eg ession coe icien s and p- alues om he Cox p opo ional haza ds eg ession model Pa ame e Reg. coe . Haza d a io SE p- alue Home ange −1.092 0.3355 0.3854 <0.01 P opo ion o ime in ese e −28.77 3.19 × 10−13 9.775 <0.01 Season o agging Sp ing 1.032 2.807 0.3754 <0.01 Home ange:P opo ion o ime in ese e 2.29 9.87 0.7639 <0.01 No e: N = 69, numbe o e en s = 38. 8 | THORBJØRNSEN ET al. (13 mm) ags educed su i al om 94% in he con ol g oup o 90% and 72%, espec i ely (Kennedy e al., 2018). Howe e , he e a e di e ences in agging p ocedu es be ween ha s udy and ou s udy, including ag posi ioning, ime om cap u e o agging and ype o seda ion agen , ha may ha e a ec ed su i al. P ese ing a spec um o di e en pe sonali ies may help sus ain a popula ion's esilience o en i onmen al change, as di e en pe sonali- ies a e a ou ed ac oss a iable en i onmen al condi ions (Dingemanse e al., 2004). In e es ingly, ish wi h di e en pe sonali ies adjus hei beha iou di e en ly when aced wi h en i onmen al change (Villegas- Ríos e al., 2018). Reac i e ish (being less bold, explo a o y and ag- g essi e han p oac i e ish) may educe hei home anges in esponse o inc easing empe a u es, while p oac i e main ain, o e en sligh ly inc ease hei s (Villegas- Ríos e al., 2018). This could u he enhance po en ial ishing- induced selec ion owa ds smalle home ange sizes o p oac i e ish. Fu he mo e, sp eading ishing e o o e a ange o habi a s could help educe cap u e bias, as di e en en i onmen- al condi ions a ou di e en beha iou s (Killen e al., 2016). Las ly, a s udy o wild- collec ed guppy popula ions showed ha ep oduc i e beha iou di e si ied in popula ions ha we e exposed o empo al he e ogenei y in p eda o biomass (Ba bosa e al., 2018). Rese es on he o he hand p o ide spa ial he e ogenei y in ‘p eda o biomass’. Ou esul s ha e clea managemen implica ions. The ac ha in- di idually consis en home ange size may a ec su i al di e en ly inside and ou side ma ine ese es implies ha , ideally, a mosaic o ma ine ese es and a eas (pa ially) open o ha es , ep esen ing a a ia ion in ishing p essu e, can p o ide a he e ogeneous selec ion egime ha can oppose di ec ional selec ion on beha iou . ACKNOWLEDGEMENTS Animal handling and agging we e conduc ed unde e hical guide- lines app o ed by he No wegian Animal Resea ch Au ho i y (Fo søksdy u alge ), pe mission # 5332 ‘Habi a use and mo emen o sea ou s udied by acous ic eleme y inside and ou side ma ine p o ec ed a eas’. Ou s udy was unded by he Resea ch Council o No way (g an No 201917 PROMAR), he FP7 ERA- Ne Biodi ERsA (g an No 225592 BUFFER), he Eu opean Union’s Ho izon 2020 esea ch and inno a ion p og amme unde he Ma ie Sklodowska- Cu ie g an ag eemen #793627 (BEMAR), he Eu opean Regional De elopmen Fund (In e eg IVa, Ma Gen II p ojec ) and he en i- onmen al o ice a he Coun y Go e no , Aus - Agde . We hank JA Knu sen and CB F ei as o ield and labo a o y assis ance. CONFLICT OF INTEREST None decla ed. DATA AVAILABILITY STATEMENT Da a a e a ailable om he D yad Digi al Reposi o y: h ps://doi. o g/10.5061/d yad.gqnk9 8sms ORCID Susanna Huneide Tho bjø nsen h ps://o cid. o g/0000-0001-7589-2339 REFERENCES Ad iaenssens, B., & Johnsson, J. I. (2011). Lea ning and con ex - speci ic explo a ion beha iou in ha che y and wild b own ou . Applied Animal Beha iou Science, 132, 90– 99. Alós, J., Palme , M., Rosselló, R., & A linghaus, R. (2016). Fas and beha io - selec i e exploi a ion o a ma ine ish a ge ed by angle s. Scien i ic Repo s, 6, 38093. Ande sen, K. H., Ma y, L., & A linghaus, R. (2018). E olu ion o boldness and li e his o y in esponse o selec i e ha es ing. Canadian Jou nal o Fishe ies and Aqua ic Science, 75, 271– 281. A linghaus, R., Laskowski, K. L., Alós, J., Kle o h, T., Monk, C. T., Nakayama, S., & Sch öde , A. (2017). Passi e gea - induced imidi y synd ome in wild ish popula ions and i s po en ial ecological and manage ial implica ions. Fish and Fishe ies, 18, 360– 373. 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