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32 (1), 2012
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Cal mo ali y o semi-domes ica ed eindee (Rangi e a andus a andus) in
he Finnish eindee -he ding a ea
Mau i Nieminen1, Ha i No be g2 & Veikko Maijala3
1 Finnish Game and Fishe ies Resea ch Ins i u e, Toi oniemen ie 246, FIN-99910 Kaamanen, Finland
(Co esponding au ho : mau i.nieminen@ k l. i).
2 Finnish Wildli e Agency, Vanamoka u 3 D, FIN-96500 Ro aniemi, Finland.
3 Uni e si y o Applied Sciences, Joki äylä 11C, FIN-96300 Ro aniemi, Finland.
Abs ac : Du ing 1999-2008 cal mo ali y was s udied in six eindee -he ding coope a i es in No he n Finland, whe e
3942 semi-domes ica ed eindee (Rangi e a andus a andus) cal es we e equipped wi h adio mo ali y colla s. The
cal es we e weighed and ea ma ked mos ly a 2-5 days o age, o a 2-8 weeks o age. Al oge he 460 dead adio-colla ed
cal es we e ound om cal ing in May un il win e ound-ups in Oc obe -Janua y. In no he n moun ain he ding
coope a i es, he a e age mo ali y o cal es a ied be ween 7-12%. On a e age, 39-54% o cal es ound dead we e
a ibu ed o p eda ion. Golden eagles killed 0-3.5% o cal es in di e en yea s and a eas in I alo and Käsi a si coope a-
i es. Golden eagles we e esponsible o 33-43% o he cases and 84-93% o all iden i ied p eda ion. Mos cal es killed
by golden eagles we e ound in July-Augus and in open a eas. Cal es killed by golden eagles we e signi ican ly (P<0.01)
ligh e han hose no p eda ed. No p eda ion occu ed in he Poikajä i coope a i e, bu he annual mo ali y o cal es
a ied be ween 0-35% in coope a i es nea he Russian bo de . In Oi anki coope a i e b own bea s killed on a e age
2% o he adio-colla ed cal es. Mos p eda ion (87%) occu ed a he end o May and in ea ly June. In he Kallioluoma
coope a i e, p eda o -killed cal es ound comp ised 53% and wol -killed 45%. P eda ion was 70% o o al mo ali y in
he Halla coope a i e, and p eda ion by wol , bea , lynx and wol e ine comp ised on a e age 38%, 20%, 9% and 2.3%,
espec i ely. The sex and pel colo did no signi ican ly a ec su i al o cal es. Bi h weigh o cal es killed by bea s was
signi ican ly (P<0.01) ligh e han hose no killed, bu hose cal es killed by lynxes we e signi ican ly (P<0.05) hea ie
han ha su i ed. Bea s killed cal es mainly in May-July, wol es in July-Oc obe and lynx in Augus -Decembe .
Key wo ds: cal mo ali y; p eda ion; adio eleme y; semi-domes ica ed eindee ; su i al.
Rangi e , 33, Special Issue No. 21, 2013: 79–90
13 h A c ic Ungula e Con e ence
Yellowkni e, Canada
22-26 Augus , 2011
In oduc ion
Se e al s udies ha e shown he majo ole o
la ge e es ial ca ni o es in neona e and ju e-
nile mo ali y o di e en ungula e species (see
Linnell e al., 1995). Mo ali y du ing he i s
summe is an impo an ac o in he popu-
la ion dynamics o many eindee (Rangi e
a andus a andus) and ca ibou (R. a andus)
he ds, and p eda ion is usually suspec ed as he
p ima y cause o mo ali y (Mille & B ough-
on, 1974; Page, 1985; Mahoney e al., 1990;
Skogland, 1991; Whi en e al., 1992; Adams
e al., 1995; Valkenbu g e al., 2004). Apa
om many he ding ac i i ies and supplemen-
a y eeding du ing win e mon hs, eindee
a e ee- anging mos o he yea . In he no h-
e n pa s o Fennoscandia semi-domes ica ed
eindee comp ise an impo an sou ce o p ey
79
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, 2012
o many p eda o s. A simila si ua ion occu s
in Finland, especially du ing he summe and
snow- ee pe iods. These a eas sha e pa s o
hei ange wi h di e en p eda o s including
wol e ine (Gulo gulo), Eu asian lynx (Lynx
lynx), b own bea (U sus a c os), wol (Canis
lupus), and golden eagle (Aquila ch ysae os).
La ge ca ni o es may cause subs an ial losses
in semi-domes ica ed eindee by p eying on
bo h adul s and ju eniles (Bjä all e al., 1990;
Nybakk e al., 2002; Danell e al., 2006; Nie-
minen, 2010). Fu he mo e, golden eagle and
ed ox (Vulpes ulpes) also p ey upon eindee
cal es (Nybakk e al. 1999; T e aa e al., 2003;
No be g e al., 2006).
The aim o his s udy was o in es iga e he
su i al, iming, ex en , and causes o eindee
cal mo ali y in six eindee -he ding coope a-
i es in he Finnish eindee husband y a ea.
S udies ha e been necessa y o assess he ole
o p eda ion in cal losses and he easibili y o
cu en compensa ion egimes o p eda o -
killed semi-domes ica ed eindee . Some esul s
a e published ea lie in Finnish epo s (No -
be g & Nieminen, 2004; 2007; No be g e al.,
2005) and in he jou nal Rangi e (Nieminen,
2010; Nieminen e al., 2011).
S udy a eas
The eindee -he ding coope a i e o I alo,
si ua ed in he municipali y o Ina i, no he n
Finland, co e s a o al land a ea o 2626 km2
(see Fig. 1). In I alo, he e a e wo he ding-
g oups, Nellim and Sou he n a ea pa ly sepa-
a ed om each o he by a ence. The eindee -
he ding coope a i e o Käsi a si, si ua ed in he
municipali y o Enon ekiö, co e s a o al land
a ea 4658 km2, and is he second la ges among
he 56 coope a i es in Finland. The coope a-
i e o Käsi a si is di ided in o h ee sepa a e
he ding-g oups by ences: 1) Palojä i, 2) Ko a
Labba, and 3) Rai ijä i. The eindee -he ding
coope a i e o Poikajä i (2414 km2) is si ua ed
in he municipali y o Ro aniemi. These s udy
coope a i es a e si ua ed in he no h bo eal
ege a ion zone (Ah i e al., 1964).
In I alo coope a i e, ange is domina ed by
olling hills wi h di e en aged o es s ands,
mainly Sco s pine (Pinus syl es is). Moun ain
bi ch (Be ula pubescens cze epanowii) g ows
on he slopes o he highes hills. In Käsi a si,
moun ains domina e he landscape. The a ea o
coni e ous o es s is ela i e small and loca ed
in he sou he n pa o he coope a i e. In Poi-
kajä i, he main landscape ypes a e Sco s pine
and No wegian sp uce (Picea abies) o es s.
Oi anki (1361 km2) and Kallioluoma (1369
km2) coope a i es a e loca ed in Kuusamo mu-
nicipali y nea he Russian bo de . The ein-
dee -he ding coope a i e o Halla in Kainuu,
also loca ed nea Russian bo de and no h o
he wild o es eindee (R. . ennicus) a ea in
Fig.1. Reindee -he ding coope a i es in he s udy and
Finnish eindee husbandy a ea: 1. I alo, 2. Käsi a si, 3.
Poikajä i, 4. Oi anki, 5. Kallioluoma and 6. Halla.
80
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Finland (municipali ies o Suomussalmi, Puo-
lanka and Hy ynsalmi), co e s a o al land a ea
3592 km2. I is he eigh h la ges coope a i e
in Finland. The coope a i es o Oi anki, Kal-
lioluoma and Halla belong o he middle bo eal
ege a ion zone (Ah i e al., 1968) (Fig. 1). The
main landscape ypes a e No way sp uce and
Sco s pine o es s wi h e icaceous hea he and
lichen and boggy a eas.
The e we e on a e age 5660 eindee (adul s
and cal es a e slaugh e ing) in he eindee -
he ding coope a i e o I alo du ing he s udy in
1999-2001, 10690 eindee in Käsi a si du ing
2002-04, 4670 in Poikajä i du ing 2001-03,
2440 in Oi anki du ing 1999-2004, and 2270
in Kallioluoma du ing 2005-06. In Oi anki,
he ac ual s udy a ea was he eas e n pa o
he coope a i e con ined o Russian bo de . In
Kallioluoma, he s udy a ea was be ween main
oad 5 and he bo de wi h Russia. The e we e
app oxima ely 1200 o 1600 adul (>1 yea
old) eindee in he coope a i e o Halla du ing
he s udy 2006-08, and yea ly abou 500-700
cal es we e bo n. Reindee densi ies (win e
s ock/km2) we e simila a 2.1 in I alo and 2.3
in Käsi a si. In Poikajä i, eindee densi y was
1.9, in Oi anki 1.8, in Kallioluoma 1.7, and
signi ican ly lowe in Halla, a 0.6-0.8 eindee /
km2.
In he en i e Finnish eindee husband y
a ea, he e we e an es ima ed >160 bea s,
>75 wol e ines, >50 lynx, and 15-25 wol es
(RKTL, 2008). Du ing 2000-03 in he en i e
no he n eindee husband y a ea (including
he 13 coope a i es) he minimum numbe o
la ge ca ni o es was: 45-50 b own bea s, 40-45
wol e ines, 5-10 lynx, and 3 wol es (Kojola
& Mää ä, 2004). The mos abundan among
he la ge ca ni o es was b own bea in I alo
and wol e ine in Käsi a si coope a i e. In he
eas e n eindee husband y a ea, he minimum
numbe o b own bea s was 80-85 and 15-20
lynx. Mos wol es o he Finnish eindee hus-
band y a ea a e ound in Kainuu (23 000 km2).
In win e 2008, he size o he wol popula ion
he e was es ima ed a 29-37 animals, a dec ease
o abou 50% compa ed wi h he p e ious yea .
The popula ion size o he lynx was es ima ed
a 140-190 indi iduals, including 23-31 li -
e s, and he size o he wol e ine popula ion a
36-53 animals (Sii a e al., 2009). Faeces sam-
ples collec ed du ing summe 2005 in Kainuu
(Näljänkä and Halla coope a i es) we e gene i-
cally analysed (DNA), and 46 unique b own
bea s we e iden i ied (RKTL, 2008). In win e
2007, 55-62 wol es we e es ima ed in wild o -
es eindee a ea o Kuhmo (RKTL, 2008).
The golden eagle popula ion in Finland is
abou 440 pai s o e i o ies. Abou 80% o
all golden eagles occu in Lapland (a mos
350 pai s in 2006; La ge Ca ni o e Wo king
G oup, 2008) and 90% in whole eindee he d-
ing a ea. In he Kainuu a ea, 11 e i o ial pai s
o golden eagles we e es ima ed in 2009 (Ollila,
2009).
Ma e ial and me hods
We i ed 3942 eindee cal es wi h mo al-
i y indica ing adio- ansmi e s (Tele il Inc.,
Lindesbe g, Sweden) ixed on expandable
neck colla s in six eindee -he ding coope a-
i es, in i e he ding-g oups and a eas du ing
1999-2008. Radio colla s weighed abou 100
g ams, abou 0.3-2.5% o he body weigh o
he cal es a ma king. Cal su i al and cause-
speci ic mo ali y we e s udied wo yea s in he
he ding g oups o Nellim (1999-2000) in I a-
lo and Rai ijä i (2002-03) in Käsi a si, and
h ee yea s in o he he ding-g oups (Sou he n
a ea o I alo 1999-2001, Palojä i and Ko a-
Labba 2002-04) in hese coope a i es and also
in coope a i es o Poikajä i, Kallioluoma and
Halla. The Oi anki coope a i e s udy con in-
ued six yea s om 1999 o 2004.
Cal es we e weighed and ma ked a 2-5 days
o age in cal ing co als in May/June. Females
we e ed in co als o 1.5 mon hs wi h silage
and concen a es du ing he sp ing and cal ing
81
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32 (1), 2012
pe iods. Mid-summe ea ma king ook place in
he las weeks o June and he i s wo weeks
o July, when cal age was 2-8 weeks. Cal es
we e ma ked in di e en co als o coope a-
i es. Cal es we e sexed and weighed, and pel
colo eco ded upon which he colla ed cal es
we e euni ed wi h hei mo he s and eleased
o summe pas u es.
Dead colla ed cal es we e loca ed by ack-
ing mo ali y senso signals om he ai ( ixed-
wing ai c a and helicop e ) and by g ound
iangula ion (senso s ac i a e a e 2.5 h s o
being mo ionless). T acking was pe o med in
2-3 day in e als du ing summe un il he end
o Augus , and once pe week in Sep embe and
Oc obe . Mo ali y ac i a ed adio-colla s we e
loca ed by using hand ecei e s (Tele il RX-
8910®, Tele il Inc., Lindesbe g, Sweden and
T acke Maxima® and hound ada s, T acke
Inc., Oulunsalo, Finland). Field obse a ions
o he si e and ca cass we e eco ded and pho-
og aphed. Cause o dea h was i s in es iga ed
in he ield (e.g., e idence suppo ing p esence
o p eda o /sca enge species, such as acks,
sca s and ea he s/downs) and hen augmen ed
by nec opsies conduc ed by biologis s in he
labo a o y o he Reindee Resea ch S a ion in
Kaamanen. The p esence o hemo haging and
pe o a ions, bo h in he skin and so issues o
he dead cal , we e c i ical o de e mining he
cause o dea h by dep eda ion. I he combined
e idence om he ield si e and he nec opsy
was inconclusi e, usually due o la e disco e y
o ca cass, he cause o dea h was classi ied as
unknown (see also Bjä all e al., 1990; No -
be g e al., 2005; 2006).
Physiological condi ion o dead eindee
cal es in Halla coope a i e was de e mined by
using he o en-d y me hod o me a a sal ma -
ow a . Condi ion was exp essed as pe cen o
ma ow a (see Nieminen & Lai inen, 1986).
S a is ical analysis
Due o he di e ence in he ma king ime and
age o cal es, he weigh o cal es a ma king
anged be ween 4.2-32 kg, and he e o e o
s a is ical analysis he weigh s we e adjus ed o
1s June and o 1s July using a daily g ow h a e
o 270 g ams (see Timisjä i e al., 1982) o
all cal es weighed in he cal ing co als in May.
A daily g ow h a e o 302 g ams was used o
emale cal es and 315 g ams o male cal es
weighed la e du ing ea ma king (No be g e
al., 2005; 2006).
The daily su i al es ima es and ´ eindee
days´ (one ´ eindee day´= one adio-colla ed
eindee ou o one day) o he adio-colla ed
cal es we e calcula ed using he Kaplan-Meie
p oduc /limi me hod (Kaplan & Meie , 1958)
and using he compu e p og am ´Kaplan-Mei-
e su i o ship analysis e sion 1.0´ (Pollock e
al., 1989) o ob ain daily and o al su i al es i-
ma es o he s udy pe iods. Daily su i al es i-
ma es we e used o p esen su i o ship cu es
be ween May/June and Oc obe . Fo calcula -
ing mon hly su i al es ima es, cause speci ic
mo ali y a es and 95% con idence limi s, he
p og am ´Mic omo e sion 1.3´ (Heisey &
Fulle , 1985) was used.
Su i al es ima es o his analysis we e cal-
cula ed based on cal es ha we e: 1) ound
dead, 2) had d opped hei adio-colla s du ing
he s udy, o 3) we e eco e ed in he au umn/
win e ound-ups (su i o s) when adio-col-
la s we e aken o . The s a is ical di e ences
in cal weigh s in di e en g oups we e es ed
using - es and s epwise logis ic eg ession. In
addi ion o weigh , he e ec o sex, pel colo ,
s udy yea , and possible in e ac ions on su i al
p obabili y we e in es iga ed using logis ic e-
g ession. S a is ical es s we e ca ied ou by use
o SPSS e . 7.0 o Windows. The da a we e
examined o s a is ical signi icance a P<0.05.
Resul s
Du ing cal ing, 460 adio-colla ed cal es we e
ound dead in he six coope a i es s udied.
Pooled su i al es ima es in he moun ain co-
82
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ope a i es o I alo (including all adio-colla ed
cal es du ing yea s 1999-2001) and o Käsi a -
si (yea s 2002-04) we e 0.937 (SE=0.011) and
0.885 (SE=0.011), espec i ely.
O 806 eindee cal es adio-colla ed in
I alo eindee -he ding coope a i e du ing
1999-2001, 4.6% (37 cal es) we e ound dead,
90.4% su i ed, and 5.0% we e no ecap u ed
un il he end o he s udy (annual moni o ing
om ma king un il he end o Oc obe ). O
919 adio-colla ed cal es in Käsi a si coope a-
i e du ing 2002-04 in o al 5.2% (48 cal es)
we e ound dead, 87.4% su i ed, and 7.4%
we e no ecap u ed. Highes a ea-speci ic an-
nual mo ali y occu ed in I alo in 2000, when
in o al 19 (51.4% o all dead cal es) we e
ound dead, and in Käsi a si in 2004, when 20
adio-colla ed cal es (41.7%) we e ound dead.
In he no he n moun ain he ding coope a-
i es o I alo and Käsi a si he a e age mo al-
i y o cal es a ied be ween 7-12% (Fig. 2).
On a e age, 39-54% o he cal es ound dead
we e a ibu ed o p eda ion, and golden eagles
killed 0-3.5% o cal es in di e en yea s and
a eas in hese coope a i es. Golden eagle p e-
da ion accoun ed o 33-43% o he cases and
84-93% o all iden i ied p eda ion. The mos
cal es killed by golden eagles we e ound du -
ing July-Augus and in he open a eas.
In I alo coope a i e, 8% o all cal es ound
dead we e killed by b own bea and 3% by
ed ox, and in Käsi a si 6% by wol e ine and
2% by uniden i ied p eda o s. Mo ali y a es
caused by p eda o s o he han golden eagle
we e on a e age less han 1% in bo h s udy co-
ope a i es. The sha e o o he iden i ied causes
o dea h (acciden s, a ic, o he s) was 19% in
bo h coope a i es, while 27-40% o dead cal es
we e associa ed wi h unknown causes o mo -
ali y. In I alo du ing 1999-2001, a o al o 8%
o all cal es ound dead we e om acciden s
and collisions wi h ehicles, while 6% we e by
acciden s in Käsi a si. O he causes (11-13%)
included disease, s ess, and poo condi ion o
cal es in bo h coope a i es. When cal es wi h
uniden i ied causes o dea h we e excluded,
p eda ion comp ised on a e age 69% o he ob-
se ed mo ali y in I alo and 74% in Käsi a si.
O 404 eindee cal es adio-colla ed in
Poikajä i eindee -he ding coope a i e du -
ing 2001-03 only 2.5% (10 cal es) we e ound
dead, 93% su i ed, and 4.5% we e no ecap-
u ed un il he end o he s udy. The a e age
mo ali y om cal ing pe iod in May o he
end o Oc obe was 5.7% and om 15 h June
o he end o Oc obe 1.4%. No p eda ion oc-
cu ed, and causes o dea h included poo con-
di ion o cal es and a ic.
Annual mo ali y o cal es a ied be ween
0-35% in coope a i es nea he Russian bo -
de . O 580 eindee cal es adio-colla ed in
he eas e n pa o Oi anki du ing 1999-2004,
7.2% (42 cal es) we e ound dead, 86.6% su -
i ed and 6.2% we e no ecap u ed un il he
end o he s udy. The a e age mo ali y om
he cal ing pe iod in May o he end o Oc-
obe was 9.7% and om 15 h June o he end
o Oc obe was 2.3%. In Oi anki, mos mo -
ali y (87%) occu ed in May and June. The
mos p ominen cause o dea h was p eda ion
by b own bea comp ising on a e age 2% o
all adio-colla ed cal es du ing 2000-04. When
cal es wi h uniden i ied causes o dea h we e
excluded, p eda ion comp ised on a e age 50%
o he obse ed mo ali y in Oi anki. F om all
iden i ied causes o dea h (n=18) 33.3% we e
killed by b own bea and wol e ine, while lynx
and wol killed 5.6%.
In he eas e n pa o Kallioluoma, 139 o
587 adio-colla ed cal es (23.7%) we e ound
dead wi hin he s udy pe iod 2005-06. Mo al-
i y o adio-colla ed cal es was on a e age 18-
19% by he end o June, and 28-29%, 36-39%
and 42-46% by he end o Oc obe , Decembe ,
and mid-Janua y, espec i ely. P eda o -killed
cal es comp ised 53% and wol -killed cal es
we e 45% o all he dead cal es ound. Wol
p eda ion was on a e age 18% while he o al
83

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a e o all p eda ion was a
leas 21%. La ge ca ni o es
comp ised 92-97% o he o al
p eda ion. The mo ali y a e
due o o he sou ces o mo -
ali y han p eda ion was on
a e age 10%. F om all iden-
i ied causes o dea h (n=100
cal es) b own bea killed 62%.
Bea and lynx killed 3%, and
golden eagle 1%.
O 546 adio-colla ed ein-
dee cal es du ing 2006-08 o-
ally 177 (32.4%) we e ound
dead du ing he esea ch pe-
iod un il mid-Janua y in
Halla coope a i e in Kainuu
a ea. The o al mo ali y in
2006-08 was 30.7% a he
end o Oc obe and inc eased
o 34.6% by mid-Janua y (see
Fig. 2). P eda ion was 70% o o al mo ali y,
and p eda ion by wol , bea , lynx, and wol e -
ine comp ised on a e age 38%, 20%, 9%, and
2.3%, espec i ely. The mo ali y o eindee
cal es was sligh ly highe in Suomussalmi nea
Russian bo de han in Hy ynsalmi/Puolanka
a ea, because o bea and wol p eda ion. The
sex and pel colo did no signi ican ly a ec
su i al o cal es.
The a e age adjus ed weigh o hose adio-
colla ed cal es ha su i ed in I alo coope a i e
was signi ican ly highe (mean 9.8 kg, SD=2.0
kg, n=169) han weigh o dead cal es (mean
7.8 kg, SD=2.2 kg, n=12; =2.79, d =179,
P=0.006). Cal es killed by golden eagle we e
signi ican ly ligh e (mean 7.2 kg, SD=2.2 kg,
n=5; =2.47, d =172) han su i ing cal es,
and also ligh e han hose cal es ha died
om o he causes (mean 8.2 kg, SD=2.4 kg,
n=7). In Ko a Labba he ding-g oup (Käsi a si
coope a i e) cal es killed by golden eagle we e
1.7 kg ligh e (mean 10.7 kg, SD=1.4 kg, n=5)
han hose cal es ha su i ed (mean 12.4 kg,
SD=1.9 kg, n=259; =1.9, d =262, P=0.053).
The a e age adjus ed weigh (on 1s July)
o hose adio-colla ed cal es killed by golden
eagles in I alo coope a i e was signi ican ly
lowe (mean 12.8 kg, SD=1.7 kg, n=11) han
mean weigh o su i o s (16.6 kg, SD=2.5
kg, n=560; =760, d =10.84, P<0.001). Also,
mean weigh o cal es killed by all p eda o s
was signi ican ly lowe (13.1 kg, SD=1.7 kg,
n=13) han mean weigh o su i ed cal es
( =7.05, d =13.17, P<0.001). The weigh o
cal es killed by golden eagle in Palojä i he d-
ing-g oup in Käsi a si was signi ican ly lowe
(mean 13.0 kg, SD=1.7 kg, n=5) han mean
weigh o su i ed cal es (19.3 kg, SD=3.4
kg, n=280; =409, d =283, P<0.001). Mean
weigh o cal es killed by all p eda o s was
signi ican ly lowe (12.2 kg, SD=2.2 kg, n=7)
han mean weigh o su i ed cal es ( =5.46,
d =285, P<0.001). Weigh o cal es killed by
golden eagle in Ko a-Labba he ding-g oup was
e y low and signi ican ly lowe (mean 7.2 kg,
SD=3.2 kg, n=6) han mean weigh o su i ed
Fig. 2. To al su i al cu es o adio-colla ed eindee cal es in I alo (in 1999-
2001), Käsi a si (in 2002-04) and Halla (in 2006-08) eindee -he ding coope -
a i es, exp essed as days a e 1 May (day 1). (Mo ali y (M) = 1 – Su i al(S)).
84
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32 (1), 2012
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cal es (13.4 kg, SD=4.3 kg, n=28; =3.03,
d =31, P=0.005). No signi ican e ec o pel
colo on cal su i al in I alo and Käsi a si co-
ope a i es was ound.
The a e age adjus ed weigh (on 1s June)
o hose adio-colla ed cal es ha we e ound
dead in Poikajä i coope a i e was sligh ly
lowe (mean 8.0 kg, SD=2.6 kg, n=10) han
mean weigh o su i o s (9.3 kg, SD=2.1 kg,
n=155; P<0.05). The a e age adjus ed weigh
o hose adio-colla ed cal es ha we e ound
dead also in Oi anki coope a i e was sligh ly
lowe (mean 9.3 kg, SD=2.5 kg, n=38) han
mean weigh o su i o s (11.0 kg, SD=2.1 kg,
n=348; P<0.001). The weigh o cal es killed
by p eda o s (8.6 kg, SD=1.2 kg, n=8) we e sig-
ni ican ly lowe han weigh o su i o s (11.0
kg, SD=2.1, n=347; =3.23, d =353, P=0.001)
(see also No be g e al., 2005). In con as o
p e ious cal mo ali y, he weigh o he cal es
in Kallioluoma coope a i e did no ha e sig-
ni ican e ec on he mo ali y.
Bi h weigh o cal es killed by bea s in Halla
was signi ican ly (P<0.01) ligh e , bu killed by
lynx signi ican ly (P<0.05) hea ie han ha o
su i o s. Bea s killed cal es mainly in May-Ju-
ly, wol es in July-Oc obe and lynx in Augus -
Decembe . Causes no associa ed wi h p eda-
ion comp ised 11.9% o o al mo ali y, and
included a ic acciden and o he acciden s.
Me a a sal a con en was < 25% in cal es
dead by bad condi ion and disease. Excluding
he dea hs om unknown causes (n=11), and
i unknown cal es ea en by di e en p eda o s,
mainly by bea s (n=18), we e also killed by
hese p eda o s, o al p eda ion was e y high,
83.1%.
Bi h weigh o cal es ha we e los o killed
by p eda o s du ing he s udy in Halla was on
a e age 0.2 kg lowe (mean 6.3 kg) han ha o
su i o s. Bi h weigh o cal es killed by bea s
was signi ican ly (P<0.05) lowe (mean 5.8
kg), bu hose killed by lynx was signi ican ly
(P<0.05) highe (mean 6.7 kg). Condi ion o
cal es was, howe e , ai o good (me a a sal
a con en > 30%). The bi h weigh o male
and emale cal es and pel colo did no a ec
su i al di e en ly, as he e we e no signi ican
in e ac ion be ween weigh , sex and colo on
cal su i al.
Discussion
Acco ding o Reime s (1983) mo ali y a es in
cal es o S alba d eindee (R. . pla y hynchus),
exis ing in an en i onmen almos ee o p ed-
a o s, a e app oxima ely 1% and 19% in he
age in e als 0-6 and 6-12 mon hs. The mo al-
i y a es among 0-6 mon h old eindee cal es
is expec ed o be wi hin he ange o 6-21%
ound also in he p eda o - ee Sou h-Geo gia
(Leade -Williams, 1980), and lowe han 45-
60% ound in many Rangi e he ds subjec o
p eda ion (Rehbinde , 1975; Be ge ud, 1980).
Wol es, bea s, and golden eagles ha e been he
mos impo an p eda o s o adio- agged ein-
dee /ca ibou cal es in bo h No h Ame ica and
Russia, as 80-89% o mo ali ies we e caused
by p eda ion (see Be ge ud, 1980).
In a s udy conduc ed in no heas e n Finn-
ish Lapland, mo ali y om golden eagles
comp ised annually 3-4% o he adio-colla ed
coho in 1997-98 (No be g e al., 2006). In
s udies conduc ed in cen al No way, Nybakk
e al. (1999) ound golden eagle p eda ion o
accoun o 1-2% among adio-colla ed cal es,
while K am e al. (1998) obse ed a o al mo -
ali y o 8%, and cal es killed by golden eagle
comp ised 40% o all cal es ound dead. In he
p esen s udy golden eagle was also he mos
signi ican cause o dea h bo h in moun ain
coope a i es o I alo and Käsi a si, causing up
o 3.5% annual mo ali y a e among adio-
colla ed cal es.
The majo i y o semi-domes ica ed eindee
cal es a e bo n in no he n Finland in May
wi h peak cal ing occu ing om 18-23 May
(Elo an a & Nieminen, 1986; Weladji e al.,
85
Rangi e , 33, Special Issue No. 21, 2013
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, 2012
2006). In his s udy, 52% o cal mo ali y in
Nellim and 87% in Ko a-Labba eindee he d-
ing g oup ook place be o e he end o June.
In Oi anki, mos mo ali y (87%) occu ed in
May-June. Acco ding o Linnell e al. (1995)
p eda ion gene ally comp ises he majo sha e
o he o al mo ali y o ju enile ungula es.
Many s udies o eindee /ca ibou ha e also
demons a ed ha cal mo ali y is usually
highes du ing he i s days and weeks a e
cal ing and hen dec eases conside ably du ing
summe and au umn (see Elo an a & Niemin-
en, 1986; Whi en e al., 1992; Adams e al.,
1995; No be g e al., 2005).
In a s udy conduc ed in cen al No way, 89%
o he o al mo ali y o cal es om Augus o
Ap il was due o p eda ion, and 60% o cal es
wi h iden i ied cause o dea h we e killed by
lynx. P eda ion comp ised an e en highe p o-
po ion, 94% o all iden i ied mo ali y, when
examined om Augus o mid-No embe (Ny-
bakk e al., 2002). Also in no he n No way,
p eda ion accoun ed o 75% o he cal losses
du ing summe and win e , and lynx was he
main p eda o (55%) (Ma hisen e al., 2003).
In Halla, p eda ion caused 70% o all cal es
ound dead and 87% o all iden i ied mo al-
i y. P eda ion by wol , bea , lynx, and wol e ine
comp ised 38.4%, 20.3%, 9.0%, and 2.3% o
all adio-colla ed cal es ound dead, espec i e-
ly (see also Nieminen, 2010).
Highes mo ali y in moun ain a eas oc-
cu ed in his s udy du ing July and Augus and
was caused mainly by golden eagles. We con-
clude ha access o, and use o alpine highlands
and o he open a eas in luenced he isk o ein-
dee cal es o p eda ion by golden eagles, and
subsequen ly he empo al su i al dis ibu ion
in s udy coope a i es. We also emphasize he
ela i e impo ance o golden eagles as a mo -
ali y ac o in he no he n pa o he Finnish
eindee husband y a ea, whe e he p opo ion
o open alpine landscape is much highe han
in he sou he n a ea (see also Nieminen e al.,
2011).
The design o he p esen s udy in Halla co-
ope a i e was simila o ha o he ou -yea
p og am ha moni o ed eindee cal mo ali y
in Sweden in he 1980s (Bjä all e al., 1990),
and also s udies in 1995-96 in cen al No way
(Nybakk e al., 2002) and in 1997-98 in no h-
eas e n Finnish Lapland (No be g e al., 2006).
The o al mo ali y eco ded in he p esen
s udy was, howe e , much highe (32.4%) han
o al mo ali y (14.3%) eco ded in Umbyn,
Sweden and in Lappi eindee -he ding coop-
e a i e in Finland (8.5%). The o al mo ali y
was also sligh ly highe han ha in No h-
T øndelag in No way (31.0%) om Augus o
Ap il.
Du ing 2006-08 in he Halla coope a i e
la ge p eda o s (mainly wol and lynx) killed
380 o 455 eindee yea ly, and compensa ion
o p eda o -killed eindee o eindee owne s
was 5-6.5 imes mo e han slaugh e incomes.
Acco ding o eindee owne s many wol es
om Russia and Kuhmo a e isi ing Halla co-
ope a i e and killing eindee mainly du ing
summe s and au umns. P eda ion accoun ed
o a highe pa o o al mo ali y eco ded
in he p esen s udy (70%) han in s udies in
Sweden (65%) and in Finland (53%). In he
No way s udy, p eda ion was highe (75.3%)
han in he p esen s udy. Indeed, ca ibou he ds
exposed o p eda ion may lose usually 50% o
he annual cal c op (Be ge ud, 1980), and p e-
da ion can cons i u e up o 93% o o al an-
nual mo ali y in cal es (Mahoney e al., 1990).
Ne e heless, he annual mo ali y eco ded in
he p esen s udy was highe han ea lie e-
po ed om Sweden and Finland. I unknown
cal es ea en by di e en p eda o s, mainly by
bea s, we e also killed by hese p eda o s, o-
al p eda ion was highes in Halla a 83%. In
cen al No way 89.3% o he o al mo ali y in
cal es was also due o p eda ion, and p eda ion
by lynx was he dominan cause (42.4%) (Ny-
bakk e al., 2002).
86
Rangi e , 33, Special Issue No. 21, 2013
32 (1), 2012
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A common pe cep ion is ha animals p eyed
upon a e ei he smalle o in poo e nu i ional
condi ion compa ed o su i o s, and se e al
s udies (e.g., Haukioja & Salo aa a, 1978;
Elo an a & Nieminen, 1986; T e aa e al.,
2003) ha e shown ha body weigh o ein-
dee cal es a cal ing and also du ing he i s
summe is posi i ely co ela ed wi h su i al.
Al hough golden eagles a e capable o killing
ungula es up o he size o an adul eindee
in ce ain condi ions (Be go, 1987), hey usu-
ally kill smalle han a e age cal es (Nybakk e
al.,1999; No be g e al., 2005; 2006). In he
adio-colla s udy conduc ed in cen al No way
(Nybakk e al., 1999), cal es we e ma ked du -
ing July and ea ly Augus , and cal es killed by
golden eagles weighed on a e age 2.7-4.1 kg
less han su i ing cal es. Also in he p esen
s udy in moun ain coope a i es, he weigh s
o cal es (adjus ed o 1s June and o 1s July)
killed by golden eagles we e 1-3.8 kg lowe
han weigh s o su i ed cal es. Mean weigh
o cal es killed by all p eda o s was also lowe
han mean weigh o cal es ha su i ed, bu
he e was no signi ican di e ence in weigh s o
p eda o -killed cal es compa ed o cal es ha
died on o he causes.
Bi h weigh o cal es ha we e los o killed
by p eda o s in Halla coope a i e was sligh ly
lowe , bu bi h weigh s o cal es killed by
b own bea s we e signi ican ly (P<0.05) lowe
han su i ing cal es. In ou ea lie s udy in
nine eindee he ding coope a i es in Finland,
bi h weigh o he los cal es was on a e age
0.4-0.5 kg lowe han bi h weigh o he su -
i ed cal es. In Oi anki coope a i e, cal es
killed by bea s also had 0.5 kg ligh e bi h
weigh compa ed o hose ha su i ed (No -
be g e al., 2002). Cal es killed by lynx in Halla
had, howe e , signi ican ly (P<0.05) highe
bi h weigh (mean 6.67 kg) han su i ing
cal es. Mos small cal es we e los o killed by
bea s du ing ea ly and mid-summe , and lynx
killed bigge eindee cal es mainly du ing au-
umn. Also in cen al No way p eda ion by
lynx peaked in au umn and ea ly win e (Ny-
bakk e al., 2002).
In he p esen s udy, golden eagles we e e-
sponsible o 33-43% o he cases and 84-93%
o all iden i ied p eda ion in moun ain coop-
e a i es in he no he n Finland. Mos cal es
killed by golden eagles we e ound du ing July-
Augus and in open a eas. P eda o -killed cal es
comp ised >50% and wol -killed cal es 45% o
all he dead cal es ound in Kallioluoma, in he
sou heas e n coope a i e nea bo de o Russia.
Ou esul s showed, howe e , highes p eda-
ion and cal mo ali y in Halla, in he sou he n
coope a i e, also nea he Russian bo de and
wild o es eindee a ea o Finland. The o al
mo ali y was >30% a he end o Oc obe and
eached 35% by mid-Janua y. P eda ion was
70% o o al mo ali y, and p eda ion by wol
comp ised on a e age 38% and by b own bea
20%. The economic consequences o eindee
husband y o his a ea makes i ques ionable o
wha ex en eindee husband y in i s p esen
o m can be con inued in Kainuu, sou heas e n
eindee -he ding egion o Finland.
Acknowledgemen s
This s udy was suppo ed inancially mainly by
he Finnish Minis y o Ag icul u e and Fo -
es y and Finnish Game and Fishe ies Resea ch
Ins i u e (RKTL). We acknowledge all he
he de s and associa es in he di e en eindee -
he ding coope a i e and he ding-g oups o o
hei suppo du ing his s udy. We especially
wan o hank he eindee owne s Viljo Hu u,
Tuomas I. Palojä i, Veikko Heiska i, Ola i
Aikkila, Heikki Hä mä, Hannu Kaa inen and
A i Jun ila. We like o hank also Pasi Koi u-
maa, Heli Rou i, Sampo Sii a and Timo Kin-
nunen and many s uden s o hei wo k in he
ield and p ojec . We also wa mly hank Sa i
and Jukka Sii a i and Heikki Tö mänen o
hei assis ance in RKTL, Reindee Resea ch
S a ion in Kaamanen.
87