E olu iona y Applica ions. 2021;00:1–9.
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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
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Re ised: 24 Feb ua y 2021
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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
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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
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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
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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.
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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
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SUPPORTING INFORMATION
Addi ional suppo ing in o ma ion may be ound online in he
Suppo ing In o ma ion sec ion.
How o ci e his a icle: Tho bjø nsen SH, Moland E, Villegas-
Ríos D, Bleeke K, Knu sen H, Olsen EM. Selec ion on ish
pe sonali y di e s be ween a no- ake ma ine ese e and
ished a eas. E ol Appl. 2021;00:1– 9. h ps://doi.o g/10.1111/
e a.13242