The
impac o
la ge
ca ni o es on he mo ali y o semi-domes ica ed ein-
dee
(Rangi e
a andus a andus
L.)
cal es
in
Kainuu,
sou heas e n eindee -
he ding egion o Finland
Mau i
Nieminen
Finnish
Game and Fishe ies Resea ch Ins i u e, Reindee Resea ch S a ion, Toi oniemen ie 246, FIN-99910
Kaamanen
([email p o ec ed]).
Abs ac .
Du ing
2006-2008
he su i al o eindee
cal es
was s udied in he eindee -he ding coope a i e o
Halla
in
Kainuu
a ea
whe e o ally 546
cal es
we e
equipped wi h adio mo ali y colla s mainly a he age o 1-3
days.
The
su i al
was moni o ed
om
he cal ing in May
un il
win e ound-ups in Oc obe o
Janua y.
The
a e,
iming and
causes
o mo ali y o eindee
we e
assessed.
In
2006-08
o ally 177 adio-colla ed
cal es
we e
ound dead (mean
mo ali y
32.4%)
un il
mid-Janua y. The
esul s
showed signi ican annual a ia ion in cal mo ali y and p eda ion.
Independen o
yea
he mo ali y o adio-colla ed
cal es
was highes du ing he i s wo mon hs a e
bi h,
and
he o al mo ali y was 30.7% a he end o Oc obe and eached 34.6% by mid-Janua y. The sex o
cal es
and pel
colou
did no a ec signi ican ly su i al o cal es. P eda ion comp ised 70.0% o o al mo ali y. P eda ion by wol ,
bea ,
lynx and wol e ine comp ised on
a e age
38.4%, 20.3%, 9.0% and 2.3%, espec i ely.
Bi h
weigh o
cal es
los
o
killed by p eda o s did no di e
om
su i ing cal es. Howe e , bi h weigh o
cal es
killed by b own
bea s
was
signi ican ly ligh e (mean 5.84 kg),
whe eas
cal es
killed by Eu asian lynx was signi ican ly hea ie (mean 6.67 kg) han
bi h
weigh o
cal es
ha su i ed (mean 6.26 kg). Bea s killed
cal es
mainly in May
o
July, wol es
in
July o Oc obe
and lynx in Augus o Decembe . O 209 adio-colla ed adul emales, 17
we e
ound dead
(8.0%).
These
emales
had
cal ed in May and hey
we e
killed mainly by wol es
(52.0%)
in Augus o Oc obe .
Key
wo ds:
cal mo ali y,
la ge
ca ni o es, p eda ion,
Rangi e ,
eindee , su i al.
In oduc ion
Se e al
s udies
ha e
shown he majo ole o
la ge
e es ial ca ni o es on he neona al
mo ali y o
la ge
and medium-sized
ungula es
(see
Linnell
e al., 1995). Reindee and ca ibou
a e no able among
ungula es
in ha hei
p o¬
ein
balance
may be
nega i e
o much o he
yea
(Ge ha e al., 1996). This may
inc ease
he impo ance o
access
o sp ing o age in
o de o
mee
high ni ogen demands
ollow¬
ing
win e consuming low-p o ein, lichen-
domina ed die s. The e o e, he p eda ion
isk- o aging ade-o may be mo e ob ious
han in o he
ungula e
species
(Gus ine e al.,
Rangi e ,
30
(1):
79
-
88
2006).
A e
ha sh win e s cal ing o semi-
domes ica ed eindee is also delayed and he
smalles
emales
die be o e cal ing o p oduce
s illbi hs.
Indeed
cal es
a e highly ulne able
o p eda o s (T e aa e
al.,
2003). Howe e , a -
e hei i s ew weeks, he p eda ion isk de-
c eases
(Adams e
al.,
1995; T e aa e
al.,
2003).
Apa
om
he ding ac i i ies and common
supplemen a y eeding du ing win e mon hs
because
o poo win e
pas u es,
semi-domes¬
ica ed eindee in Finland a e ee- anging o
mos o he
yea ,
especially so du ing he sum-
me season. Supplemen a y eeding has a
posi¬
i e e ec on sp ing body weigh especially o
Rangi e ,
30
(1),
2010
79
Fig.
1. S udy
a ea,
he eindee -he ding coope a i e o
Halla,
whe e eindee cal es and also adul emales
we e i ed
wi h
adio-colla s in di e en co als (•) in 2006-08. The illage o
Hy ynsalmi
is shown (*).
smalle eindee emales (Ba dsen e
al.,
2008).
The
condi ion
o supplemen a y ed eindee
is usually good du ing win e and sp ing, and
cal -p oduc ion
and
bi h
weigh o newbo n
cal es a e also good (Soppela &
Nieminen,
2001).
Howe e ,
locally
la ge
ca ni o es may
cause
signi ican
losses
o eindee husband y. Ac-
co ding
o eindee he de s in
Finland,
cal
mo ali y
is ex ensi e be ween cal ing,
mid-
summe
ea -ma king ound-ups and subse-
quen au umn and win e ound-ups in
Oc o-
be o
Janua y.
B own
bea
U sus
a c os,
wol
Canis
lupus,
Eu asian lynx Lynx
lynx,
wol e ine
Gulo
gulo
and golden
eagle
Aquila
ch ysae os
a e
all
po en ial p eda o s in
Finnish
eindee -
he ding
coope a i es
(No be g
e al., 2005:
No be g
&
Nieminen,
2007).
The
objec i e o his s udy was o examine
he mo ali y among eindee , cal es
om
he
age o 1-3
days
o 7-8 mon hs and adul e-
males
in
Halla
eindee -he ding coope a i e
in
Kainuu
a ea
nea he Russian bo de and
abou 20 km o he
no h
o he
wild
o es
eindee
(Rangi e
a andus
ennicus
Lonnb.)
a ea.
S udy
a ea
The
8 h
la ges
eindee -he ding coope a i e
in
Finland
Halla
wi h
70 eindee he de s is
si ua ed in he municipali ies o Suomussalmi,
Puolanka
and
Hy ynsalmi
(Hy ynsalmi
illage,
64o40'N,
28o29'E,
Fig. 1) and i co e s a o al
land
a ea
3592 km2 in he
Kainuu
a ea,
which
belongs o he middle bo eal ege a ion zone
(Ah i
e
al.,
1968). The landscape is domina ed
by
No way
sp uce
Picea
abies
and Sco ch pine
Pinus
syl es is
o es s
wi h
e icaceous hea he ,
lichen
and boggy
a eas.
The e we e 1200 o
1600 adul (>1
yea
old) eindee in he co¬
ope a i e du ing he s udy, and abou 500-700
cal es we e
bo n
annually.
80 Rangi e ,
30
(1),
2010
Ma e ial and
me hods
The
s udy was ca ied ou du ing he
yea s
2006 o 2008. The emales we e ounded up
in
cal ing co als and ed o 1.5
mon h
wi h
silage
and concen a es
(Po onHe kku,
Rehu-
Raisio)
du ing he sp ing and cal ing
pe iod.
Cal es
we e weigh ed and o ally 546 cal es
(286 males, 260 emales) (106 cal es in 2006,
260 in 2007 and 180 in
2008)
equipped
wi h
silen
mo ali y de ec ing adio- ansmi e s
(Tele il
Inc., Lindesbe g, Sweden; equen-
cies 138
MHz
and 230
MHz)
ixed
on expand¬
able neck colla s.
This
was mainly done a he
ea ly age o one o h ee days.
Ma king
ook
place yea ly also in he las weeks o June and
he i s weeks o July, when o ally 78 o he
cal es we e adio-colla ed a he age o
ou
o
six weeks. The colla s weighed abou 100
g ams and co esponded o 0.7-2.3% o he
body
weigh o he cal es a he s a .
Mos
o
he cal es we e ma ked in he Suomussalmi
a ea
(72.9%).
Each
cal
was sexed and pel
colou
was eco ded on a i e-s ep scale (1 =
whi e,
2 = ligh , 3 =
no mal
b own,
4 = da k
b own
and 5 = black). In addi ion, 209 adul
emales (100 emales in 2007, 109 (10 wi hou
cal es)
in 2008, abou 8% o all emales) we e
equipped
wi h
mo ali y adio- ansmi e s.
A e
ma king, he animals we e adio- acked
by
g ound
iangula ion o loca e dead animals.
T ackings
we e pe o med in in e als o wo
o
h ee
days
un il
he end o Sep embe .
Du -
ing
Oc obe
o Janua y, g ound iangula ions
we e made once pe week.
A e
loca ing
he ac i a ed ansmi e s,
pe ¬
sons
ound
hem using
hand
ecei e s (Tele il
RX-8910®,
Tele il Inc., Lindesbe g, Sweden
and
hound
ada
modi ica ion
by T acke Inc.,
Oulunsalo,
Finland).
Cause o dea h was i s
in es iga ed in he
ield
(e.g. e idence sup-
po ing
p esence o p eda o /sca enge spe-
cies, such as acks,
sca s
and ea he s/downs)
and
hen suppo ed by nec opsies conduc ed
by
biologis s in he labo a o y a he Reindee
Resea ch S a ion in Kaamanen. The p esence
o
haemo hages 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
when
dep eda ion was suspec ed (see Bja all,
e al. 1990). I he combined e idence
om
he
ield
si e and he nec opsy was inadequa e,
usually due o
la e
disco e y o ca cass, he
cause
o dea h was classi ied as
unknown.
The
s udy pe iods we e e mina ed in
la e
Janua y
o
he
ollowing
yea s,
when he majo i y o
cal es and some adul eindee we e selec ed
o
slaugh e du ing he ound-ups in au umn
and
win e .
Physiological
condi ion
o dead eindee
was de e mined by using he o en-d y
me hod
o
me a a sal ma ow a .
Condi ion
was ex¬
p essed as pe cen o ma ow a (see
Niemin-
en
&
Lai inen,
1986).
S a is ical
analysis
The
daily su i al es ima es and " eindee
days"(one " eindee day"= one adio
col¬
la ed eindee ou o one day) o he adio-
colla ed
cal es we e calcula ed using he Ka-
plan-Meie
p oduc /limi me hod
(Kaplan
&
Meie ,
1958) and using he compu e p og am
"Kaplan-Meie
su i o ship analysis e sion
1.0"
(Pollock
e ail, 1989) o ob ain daily and
o al
su i al es ima es o he s udy pe iods.
Daily
su i al es ima es we e used o p esen
su i o ship
cu es be ween May and Janua y
o
he
yea s
2006-08.
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. Di e ences in mon hly su ¬
i al
a es
be ween he h ee
yea s
we e es ed
using
ANOVA
mul i a ia e analysis o
a i¬
ance. The su i al es ima es we e calcula ed
based on documen ed
cases,
i.e. in he su i al
analysis
including
only he cal es ha 1) we e
ound
dead, 2) had d opped hei adio-colla s
du ing
he s udy o 3) we e eco e ed in he
Rangi e ,
30
(1),
2010
81
Table
1.
Mo ali y
o
adio-colla ed
cal es (n=546) in di e en
a eas
in
Halla
eindee -he ding
coope a i e
in
2006-08.
Suomussalmi
Hy ynsalmi/
Puolanka
To al To al
Dea h
causes
2006 2007 2008 2006 2007 2008 2006 2007 2008
2006¬
08
Wol e ine
1
3
1
3
4
Lynx
1 8 7 8 8 16
Wol
10
22
15 7 10 4 17
32
19 68
B own
bea
2
32
2 2
32
2
36
T a ic
1
2
1
3
1 4
D owning
3 3 3
Knocking
by o he
2
1
2
1
3
emales
2
1
2
1
3
Colla
acciden 1 1
2 2
Bad
condi ion
1 4 1 4 5
Disease
1 1 1 1
2
Unknown
3
1 5 1 1 4 1 6 11
Unknown,
ea en by 1 1 1
wol e ine
1 1 1
Unknown,
ea en by lynx 1 1 1
Unknown,
ea en by
wol
2
1
2
1
3
Unknown,
ea en by 4 8 5 1 4 9 5 18
b own
bea 4 8 5 1 4 9 5 18
Radio-colla ed
cal es,
o al
55 200 143 51 60 37 106 260 180 546
Dead
cal es,
o al
25 75 33 20 18 6 45 93 39 177
Dead
cal es, % 45.5 37.5 23.1 39.2 30.0 16.2 42.5 35.8 21.7 32.4
win e
ound-ups (su i o s) when
adio-col¬
la s we e aken o . All o he s a is ical
es s
(bi h
weigh , sex, pel
colou ,
s udy yea and
possible
in e ac ions) 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
O
546 adio-colla ed eindee cal es
du ing
2006 o 08, a
o al
o 177 (92 males, 85 e¬
males) we e
ound
dead be ween
ma king
in
May
o June/July and mid-Janua y (Table 1).
The
mean
cal
mo ali y was 32.4% in
Hal¬
la
eindee -he ding coope a i e, and i was
sligh ly
highe in
Suomussalmi
(33.4%)
han in
Hy ynsalmi/Puolanka
a ea (29.7%). The
mo ¬
ali y
was highes
(42.5%)
in 2006 and lowes
(21.7%)
in 2008. The e we e g ea annual
di ¬
e ences in
su i al
o cal es
du ing
he s udy
pe iod
(P<0.001).
Be ween
1
May
and
15
Janu¬
a y,
su i al
o
adio-colla ed
cal es was 0.265
(SE
=
0.080)
in he 2006
coho ,
0.580 (SE
=
0.030)
in he 2007
coho
and 0.711 (SE =
0.036)
in he 2008
coho .
To al
su i al
cu e
o
cal es o he 2006 o 2008
coho
was
0.654 (SE
=
0.036)
(Fig.
2).
The
mo ali y (M=1-Su i al) o
adio-
colla ed
cal es was g ea
du ing
he
i s
wo
82
Rangi e ,
30
(1),
2010
mon hs
a e
bi h and eached
31.7% by he end o
June
in 2006,
21.1% in
2007
and 13.4% in 2008.
Mo ali y
o
cal es
was 46.4%
a he end o Augus in 2006,
59.9% a he end o Oc obe and
eached 73.5% by mid-Janua y. In
2007
mo ali y o adio-colla ed
cal es
was 37.9% a he end o
Oc obe
and 42.0% in mid-Jan¬
ua y.
In
2008
mo ali y o
cal es
was 28.9% a he end o Oc obe
and
also
28.9% in mid-Janua y.
The
o al mo ali y in
2006-08
was 30.7% a he end o Oc obe
and eached 34.6% by mid-Janu¬
a y (see Fig. 2). The bi h weigh
o
male
and
emale
cal es
and
pel colou did no signi ican ly a ec su i al
di e en ly, as
he e
we e
no signi ican in e ac¬
ion
be ween sex and colou on cal su i al.
P eda ion comp ised 70.0% o o al mo ¬
ali y (p eda ion ou o o al numbe o
dead
adio-colla ed
cal es
eco e ed)
du ing
h ee
yea s
in
2006
o 2008. P eda ion showed sig¬
ni ican
(P<0.001)
annual a ia ion. P eda ion
by
wol
comp ised on
a e age
38.4%, by
bea
20.3% and by lynx 9.0% o all adio-colla ed
cal es
(Fig. 3). Wol e ines killed only ou
ma ked
cal es
(2.3 %) du ing his
s udy,
one in
May
and
h ee
in
Janua y.
Bea s
killed eindee
cal es
mainly in May-July, wol es
in
July-Oc o¬
be and lynx in Augus -Decembe . No
cal es
we e
killed by golden
eagles
in he coope a i e
o
Halla.
Causes
no
associa ed
wi h p eda ion com¬
p ised 10.7% o o al mo ali y, and included
a ic
acciden s
(4) and o he
acciden s
(8)
and o he known
causes
(7). Me a a sal a
con en was <25% in
cal es
ha
died
om
s a a ion (5) and
disease
(2). Excluding he
dea hs
wi h unknown
causes
(11), and i he
o he unknown
dea hs
o
cal es
ea en
by di ¬
e en p eda o s (see Fig. 3), mainly by
bea s
Fig.
2. Su i o ship
cu es
o adio-colla ed eindee
cal es
in
Halla eindee -he ding coope a i e in 2006-08,
ela ed
o ex-
p essed
as
days
a e
1 May (day 1). To al su i al cu e is S( )
2006-2008. (Mo ali y (M) = 1 - Su i al (S)).
Fig.
3. Dea h
causes
o
cal es
in Halla eindee -
he ding coope a i e in 2006-08.
(18),
we e
also
caused
by
hese
p eda o s, o al
p eda ion was e y high, 83.1%.
Bi h
weigh o all
cal es
ha
we e
los o
killed
by p eda o s du ing he s udy was on a ¬
e age
0.2 kg, bu no signi ican ly lowe han
ha
o su i ed
cal es
(mean
6.26 kg).
Bi h
weigh o
cal es
killed by wol es was no sig¬
ni ican ly bu by
bea s
signi ican ly (P<0.05)
lowe
(mean
5.84 kg) and by lynx signi ican -
Rangi e ,
30
(1),
2010
83

ly
(P<0.05) highe (mean 6.67 kg) han
bi h
weigh o su i ing cal es.
Condi ion
o all
p eda o killed
cal es
was, howe e , ai o
good
(me a a sal
a con en >30%).
To ally
17 adio-colla ed adul
emales
(8%)
we e
ound
dead du ing 2007-2008
un il
mid-
Janua y;
o hose nine
we e
killed by wol es
(52.0%).
Females
we e
killed mainly du ing
Augus
o Oc obe , and hey
we e
in good
condi ion
(me a a sal
a con en >60%). All
dead
emales
had cal ed in May.
Discussion
The
design o he p esen s udy was a he
simila
o ha o he eindee cal mo ali y
s udy in Sweden in he
1980s
(Bja all e al,
1990), o he
s udies
in 1995-1996 in mid
No -
way (Nybakk e al,
2002)
and in 1997-1998 in
no heas e n
Finnish
Lapland (No be g e al,
2006). Howe e , he o al mo ali y eco ded
in
he p esen s udy was much highe
(32.4%)
han he o al mo ali y
(14.3%)
eco ded in
Umbyn,
Sweden and in
Lappi
eindee -he d¬
ing
coope a i e in
Finland
(8.5%)
bu simila
o
ha in No h-T ondelag in mid No way
(31.0%).
P eda ion
accoun ed o a
la ge
pa o o al
mo ali y
eco ded
in
he p esen s udy
(70.0%)
han in
s udies
in Sweden (65%) (Bja all e al,
1990) and in
Finland
(53%) (No be g e al,
2006). In he No wegian s udy, he pa o
p eda ion
was e en highe
(75.3%)
(Nybakk e
al,
2002)
han in he p esen s udy.
Acco ding
o
Be ge ud
(1980)
ca ibou he ds exposed o
p eda ion
may lose usually 50% o he annual
cal
c op, and p eda ion has been epo ed o
cons i u e up o 93% o he o al annual mo ¬
ali y in
cal es
(see Mahoney e al, 1990). I he
o he unknown dea h
causes
o
cal es
ea en
by
di e en p eda o s, mainly by
bea s,
we e
eally killed by
hese
p eda o s, o al p eda ion
was highes in p esen s udy, o ally 83.1%. In
ou
s udy
om
Kuusamo, also in he sou h¬
eas e n
eindee -he ding egion in
Finland,
he
o al
mo ali y o eindee
cal es
was 42-46%
by
mid-Janua y in 2005-2006, and he o al
a e
due p eda ion was a
leas
21%.
P eda o -killed
cal es
comp ised 53% o all o he dead
cal es
ound
(No be g
&
Nieminen,
2007).
Mainly
wol es and
bea s
caused
g ea
losses
among
cal es
in he p esen s udy, and p eda¬
ion
by
wol
comp ised on
a e age
as much as
38.4% o he adio-colla ed cal es. Bea s killed
cal es
mainly
om
May o July, wol es in July
o
Oc obe and lynx in Augus o Decembe .
Also
in
Kallioluoma
eindee -he ding
coope a¬
i e in Kuusamo
a ea,
mos o he dead
cal es
ound
(53%)
we e
killed by p eda o s and o
hem
45% by wol es.
Wol
p eda ion was on
a e age
18% while he o al
a e
o p eda ion
was a
leas
21%
(No be g
&
Nieminen,
2007).
In
mid No way 89.3% o he o al mo ali y in
cal es
was due o p eda ion, and p eda ion by
lynx
was he dominan
cause
(42.4%)
(Nybakk
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 he lynx was he
main
p eda o (55%) (Ma hisen e al, 2003). In
no heas e n
Finnish
Lapland a minimum o
53%
o he o al mo ali y was a ibu ed o
p eda ion,
wi h golden
eagles
being he main
p eda o . Mo ali y caused by golden
eagles
comp ised,
howe e , only 2.8% o 4.2% in
1997-99 (No be g e al, 2006). In an ea lie
s udy in mid No way also 5.3 % o he o¬
al
loss o adio-colla ed eindee
cal es
was
caused by golden
eagles
(Nybakk e al, 1999).
The
golden
eagle
popula ion in whole
Fin¬
land
is a ound 440 pai s o e i o ies.
Abou
80% o all golden
eagles
a e
li ing
in Lapland
(maximally 350 pai s in 2006, La ge Ca ni o e
Wo king
G oup,
2008)
and 90% in he whole
eindee -he ding
a ea.
In
Kainuu
a ea,
he e
we e
only 11 e i o ial pai s o golden
eagle
in
2009
(Ollila,
2009). In he p esen s udy,
gold¬
en
eagles
killed no eindee
cal es
and adul
emales
in he o es
a ea.
In mo ali y
s udies
conduc ed
in mid No way and in no heas -
84 Rangi e ,
30
(1),
2010
e n
Finnish
Lapland,
he mos o he adio-
colla ed
eindee
killed
by golden
eagles
we e
disco e ed
in
open
alpine e ain (see
Nybakk
e al, 1999;
No be g
e al, 2006). Inc eased isk
o
golden
eagle
p eda ion
seems
o associa e
wi h
alpine highlands.
Bi h
weigh s o cal es ha we e los o
killed
by p eda o s in he p esen s udy was
sligh ly
lowe ,
bu cal es
killed
by
b own
bea s
signi ican ly
(P<0.05) lowe , han
bi h
weigh
o
he cal es ha
su i ed.
In an ea lie s udy
in
nine eindee he ding coope a i es in
Fin¬
land,
bi h
weigh s o he los cal es was on
a e age
0.4-0.5 kg lowe han
bi h
weigh o
he
su i ing
cal es. In
Oi anki
coope a i e in
Kuusamo
a ea, cal es
killed
by bea s we e 0.5
kg
ligh e
a
bi h
compa ed o hose
su i ing
(No be g
e al, 2002). Cal es
killed
by lynx in
his s udy
had,
howe e , 0.4 kg and signi ican ¬
ly
(P< 0.05) highe
bi h
weigh s (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.
In mid
No way,
p eda ion
by lynx also peaked in au umn and
ea ly win e
(Nybakk
e al, 2002).
In
he p esen s udy he
bi h
weigh o male
and
emale cal es and pel
colou
did no a ¬
ec
su i al.
The e was also no signi ican
in¬
e ac ion
be ween sex and pel
colou
on
cal
su i al.
In s udies on ungula es whe e bias
in
neona al p eda ion is appa en , i is usually
male biased, and
acco ding
o
Aanes &
Ande -
son
(1996)
sex di e ence in
cal
beha iou
has
been
sugges ed as po en ial explana ion.
Ac¬
co ding
o
Ma hisen
e
al.
(2003)
male eindee
cal es s ay usually a he away
om
hei
mo he s,
exhibi a highe le el o
locomo i e
beha iou
in e ms o play and walking, and
a e he e o e mo e ulne able o p eda ion
han
a e emales cal es.
Du ing
he las
i e
yea s
p eda o s ha e
caused inc eased losses in he
Finnish
semi-
domes ica ed
eindee s ock ( o ally 3500-4000
eindee /yea ), especially in he sou heas e n
egions,
and mainly due o he expansion o
he
wol
popula ion
in o
he
Kainuu
a ea.
Based
on eindee he ding s a is ic
om
1976
o
2008 la ge ca ni o es ha e
killed
a ound
2000 eindee (based on eindee
ound
dead)
annually in he
Finnish
eindee -he ding a ea.
The
eindee we e
killed
mos ly by wol e ines
(a ound
45%),
b own
bea (25%),
wol
(20%)
and
lynx (10%)
(Nieminen,
2009).
Du ing
1994-2008 la ge ca ni o es ha e
killed
mainly
adul
emales (59.2%), cal es (<1 yea -old)
(27.4%)
and adul males (9.5%). The cal es
we e
killed
mainly by bea s (a ound 30%),
wol¬
e ines (29%), wol es (23%) and lynx (18%)
(Nieminen,
2009).
Bea
sca s
collec ed
du ing
summe 2005
in
whole
Kainuu
(23 000 km2), also ou side
he eindee husband y a ea, we e analysed
using
aecal
DNA-me hodology,
and a o al
o
46
b own
bea s we e inden i ied
(RKTL,
2008), and in win e 2007, 55-62 wol es we e
es ima ed o li e in he nea by
Kuhmo
a ea.
In
he whole
Finnish
eindee husband y
a ea, he es ima ed
minimum
numbe s o he
p eda o s we e es ima ed 15 o 25 wol es and
mo e
han160 bea s,
75
wol e ines
and 50 lynx
(RKTL,
2008). The majo i y o he oday
wol
popula ion
in he eindee husband y a ea is
ound
in
Kainuu,
in he
a eas
o he
Halla
and
Naljanka eindee -he ding coope a i es.
In
win e 2008 he size o he
Kainuu wol
popula ion
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,
includ¬
ing
23—31
li e s, and he size o he wol e ine
popula ion
a
36—53
animals (Sii a e al, 2009).
Du ing
2006 o 2008 big p eda o s (mainly
wol
and lynx)
killed
yea ly 380 o 455
ein¬
dee in
Halla,
and compensa ions o p eda o
killed
eindee o eindee owne s we e 5-6.5
imes mo e han slaugh e incomes.
Thus,
many
wol es
om
Russia and
om
he
wild
Rangi e ,
30 (1), 2010
85
o es eindee
a ea
Kuhmo
a e isi ing
Halla
eindee -he ding coope a i e
killing
eindee
mainly du ing summe and au umn mon hs
(Niem nen,
2009).
My
esul s
showed e y high p eda ion and
cal
mo ali y in
Halla
eindee -he ding
coop¬
e a i e, and he economic consequences
makes
i
ques ionable o wha ex en eindee hus-
band y in i s p esen
o m
can be con inued
in
Kainuu,
he sou heas e n eindee -he ding
egion
o
Finland.
Acknowledgemen s
This
s udy was suppo ed inancially mainly by he
Finn-
ish
Minis y
o Ag icul u e and Fo es y and
Finnish
Game
and Fishe ies Resea ch Ins i u e. I acknowledge
all
he he de s and
associa es
in he eindee he ding
co¬
ope a i e o
Halla
o hei suppo du ing his s udy. I
wan o hank he eindee owne s
Hannu
Kaa inen and
A i
Jun ila, and also Pen i
Heikkinen,
Mikko
Jun unen,
Heikki
Kemppainen,
Ma kku
Ma e o,
Au o
Me inen,
Asko
Moilanen,
Ossi
Pyykkonen,
Ola i
Pollanen,
Usko
Seppanen,
Esko
Toi anen,
Ola i
Tol anen and
Ola i
Vaisanen
o hei pa icipa ion and con idence in my
wo k.
I especially wan o hank
M .
Timo
Kinnunen
and
Sa u Tol anen o hei wo k in he
ield,
and
Ha i
No -
be g, Sa i and Jukka Sii a i and
Heikki
To manen a he
RKTL,
Reindee Resea ch S a ion in Kaamanen o hei
assis ance.
I like o hank
Øys ein
Holand
o his aluable
commen s o my manusc ip . The anonymous e iewe s
a e also acknowledged o hei commen s.
Re e ences
Aanes, R. & Ande sen, R. 1996. The e ec o sex, ime
o
bi h,
and habi a on he ulne abili y o oe
dee
awns o ed ox p eda ion.
—
Can. J.
Zool
74: 1857¬
1855.
Adams,
L.G.,
Sp inge ,
F.J., & Dale, B.W 1995.
Ca i-
bou
cal mo ali y in
Denali
Na ional
Pa k, Alaska.
—
Jou nal
o
Wildli e
Managemen
59(3):
584-594.
Ah i,
T., Hame -Ah i, L., &
Jalas,
J. 1968. Vege a ion
zones and hei sec ions in no hwes e n
Eu ope.
— An¬
nales
Bo anica
Fennica
5: 169-211.
Be ge ud,
A.T.
1980. A e iew
o
he popula ion dynam¬
ics o ca ibou and
wild
eindee in
No h
Ame ica.
—
In: Reime s, E., Gaa e, E. & Skjennebe g, S.
(eds.):
Second
in e na ional
eindee /ca ibou
symposium.
Di ec-
o a e
o
il
og
e sk ann isk,
T ondheim,
No way, pp.
556-581.
Bja all,
A.,
F anzén,
R., No dk is , M., &
Åhman,
G.
1990. Rena och o dju .
—
Na u d d e ke
o lag,
Solna,
Sweden, 296pp. (In Swedish).
Ba dsen, B.-J., Fauchald, P., T e aa, T. Langeland, K.,
&
Nieminen,
M.
2008.
Expe imen al
e idence
o
cos s
o
lac a ion in a low isk en i onmen o a long-li ed
mammal.
-
Oikos
118:837-852.
Ge ha ,
K.L.,
Whi e,
R.G., Came on,
R.D.,
& Russell,
D.E.
1996.
Body
composi ion
and nu ien
ese es
o
a c ic ca ibou.
—
Canadian
Jou nal
o
Zoology
74: 136¬
146.
Gus ine,
D.G.,
Pa ke ,
K.L.,
Lay, R.J., Gillingham, M.
P.,
& Hea d, D.C. 2006.
Cal
su i al o woodland
ca ibou
in a mul i-p eda o ecosys em.
—
Wildli e
Mon-
og aphs
ol.
165 (1): 1-32.
Heisey,
D.M.
& Fulle ,
T.K.
1985.
E alua ion
o
su i al
and
cause
speci ic mo ali y
a es
using eleme y da a.
—Jou nal
o Wildli e
Managemen
49(3):
668-674.
Kaplan,
E.L.
& Meie , P. 1958. Nonpa ame ic es ima-
ion
om
incomple e obse a ions.
—
Jou nal
o he
Ame ican
S a is ical
Associa ion
53: 457-481.
La ge
Ca ni o e
Wo king G oup 2008.
—
In: Ekens ed ,
J.
& Schneide , M.
(eds.).
The
golden
eagle
(Aquila
ch ysae osj in he
No h
Calo e
A ea
1990-2007.
The
No h
Calo e
Council,
Repo 55, 45pp.
Linnell,
J.D.C.,
Aanes, R., & Ande sen, R. 1995. Who
killed
Bambi? The ole o p eda ion in he neona al
mo ali y
o empe a e ungula es.
—
Wildli e
Biology
1(4): 209-223.
Mahoney, S.P., Abbo ,
L.H.,
Russell, H., &
Po e ,
B.R.
1990.
Woodland
ca ibou cal mo ali y in insula
New oundland.
—
In: My be ge
(ed.).
T ansac ions
o
he
XlX h
cong ess
o
In e na ional
Union
o
Game
Biolo-
gis s,
T ondheim,
No way,
Vol.
I,
Wildli e
Managemen ,
pp.
592-599.
Maijala,
V.,
No be g,
V.,
Kumpula, J., & Nieminen, M.
2002.
Po on
asa uo o ja -kuolema Suomen
po on-
hoi oalueella. (In
Finnish
wi h an
English
summa y:
Cal
p oduc ion
and mo ali y in he
Finnish
eindee
he ding
a ea).
—
Kala-
ja
Riis a apo eja
n o 252: 1-61.
Ma hisen,
J.H.,
Landa, A., Ande sen, R., & Fox, J.L.
2003. Sex-speci ic di e ences in eindee cal beha -
iou
and p eda ion ulne abili y.
—
Beha io al
Ecology
14(1):
10-15.
Nieminen,
M. 2009. The impac o
la ge
ca ni o es on
he mo ali y
o
semi-domes ica ed eindee in
Kainuu,
sou heas e n eindee -he ding egion o
Finland.
—
In:
XXIX
In e na ional
Union
o
Game
Biologis s
Cong ess,
Augus
17-22,
2009,
Moscow,
pp. 236-237.
Nieminen,
M. & Lai inen, M. 1986. Bone ma ow and
kidney a as indica o s o condi ion in eindee .
—
Rangi e ,
Special
Issue
1: 219-226.
No be g,
H.
& Nieminen, M. 2007. Suu pe ojen aiku-
us po on asojen kuolleisuu een
Kallioluoman
palis-
kunnassa uosina 2005-06. (In
Finnish
wi h an Eng-
86 Rangi e ,
30 (1), 2010
Pollock,
K.H.,
Wm e s ein,
S.R., & Con oy, M.J.
1989.
Es ima ion
and
analysis
o
su i al
dis ibu ion
o
adio
agged
animals.
—
Biome ics
45: 99-109.
RKTL
2008.
Kuhmos a
loy yi
DNA-analyysissa
46 ka -
hua.
Suomalais en ka hujen ulos eis a on ensimmais a
ke aa eh y
DNA-
eli pe in o ekijaanalyysi. h p://
www. k l. i/ iedo ee /kuhmos a
(In
Finnish).
Sii a, A., Ke anen, J., & Kojola, I. 2009.
Kainuun
su-
u pe ojen lumijalkilasken a 2008 (In
Finnish
wi h
an
English
abs ac : La ge ca ni o es snow- ack
coun ing
in
Kainuu
2008).
—
Riis a-
ja
kala alous
—
Tu kimuksia
1/2009.
30pp.
Soppela,
P. &. Nieminen, M. 2001. The e ec o
win¬
e ime unde nu i ion on he a y acid
composi ion
o
leg bone ma ow
a s
in eindee
(Rangi e
a andus
a andus
L.).
—
Comp.
Biochem.
Physiol.
Pa B 128:
63-72.
T e aa, T., Fauchald, P., Henaug, C., &
Yoccoz,
N.G.
2003. An examina ion
o
a
compensa o y ela ionship
be ween
ood
limina ion and p eda ion in semi-do¬
mes ic eindee .
—
Oecologia
137: 370-376.
Manusc ip
ecei ed
27
May,
2010,
e isions
accep ed
14
Decembe ,
2010
Suu pe ojen aiku us po on asojen
(Rangi e
a andus a andus
L.)
kuolleisuu een
Kainuussa,
Suomen
po onhoi oalueen
kaakkoisosassa
Abs ac
in
Finnish
/
Tii is elma.
Vuosina
2006-2008 u ki iin po on asojen sel iy ymis a
Kainuun
alueella
Hallan
pal-
iskunnassa, jossa yh eensa 546
asalle
lai e iin kaulaan kuole uuslahe ime
paaasiassa
1-3 uo okauden
iassa.
Vaso-
jen sel iy ymis a seu a iin oukokuises a asonnas a al ie o uksiin saakka loka- ammikuun aikana. Tu kimuksessa
sel i e iin
po ojen kuolemien suu uu a, ajoi umis a ja syi a.
Vuosina
2006-08 loyde iin kuolleena yh eensa 177 a-
diopan a asaa (keskimaa ainen kuolleisuus
32,4%)
ammikuun
puoli aliin
mennessa. Tulokse osoi i a me ki se aa
uosi ais a asakuolemien ja pe ojen osuuden aih elua. Riippuma a uosis a adiopan a asojen kuolleisuus oli su-
u in a
kahden ensimmaisen kuukauden aikana asonnas a, ja kokonaiskuolleisuus oli 30,7%
lokakuun
lopussa ja 34,6%
ammikuun
puoli alissa. Vasojen sukupuolella ja
u kin
a illa ei ollu me ki se aa aiku us a asojen sel iy ymiseen.
Pe ojen
osuus oli 70,0% kokonaiskuolleisuudes a. Suden, ka hun, il eksen ja ahman osuude kuolleisuudes a oli a as-
aa as i keskimaa in 38,4%, 20,3%, 9,0% ja 2,3%. Mene e yjen ai ape ujen asojen syn ymapaino ei a e onnee sel-
iy yneiden
asojen syn ymapainois a.
Ka hun
appamisen asojen syn ymapaino
oli
kui enkin
me ki se as i pienempi
(keskia o 5,84 kg) mu a il eksen appamien asojen me ki se as i suu empi (keskia o 6,67 kg)
kuin
sel iy yneiden
asojen syn ymapaino (keskia o 6,26 kg).
Ka hu
appoi a asoja
paaasiassa
ouko-heinakuun,
sude heina-lokakuun
ja il ekse elo-joulukuun aikana. 209 adiopan a aa imes a loyde iin kuolleena 17
(8,0%).
Nama
aa ime oli a a-
sonee oukokuussa, ja
paaasiassa
sude
(52,0%)
oli a
appanee
ne elo-lokakuun aikana.
lish
summa y: The impac o
la ge
ca ni o es on he
mo ali y
o semidomes ica ed eindee
cal es
in he
eindee -he ding coope a i e o
Kallioluoma
in 2005¬
06). -
Kala-
ja
Riis a apo eja
n o 415: 1-55.
No be g,
H.,
Kojola, I., Aikio, P., &
Nylund,
M. 2006.
P eda ion
by golden
eagle
Aquila
ch yae os
on semi-do-
mes ica ed eindee
Rangi e
a andus
cal es
in
no h-
eas e n
Finnish
Lapland.
-
Wildli e
Biology
12(4):
423¬
432.
Nybakk,
K.,
Kjel ik,
O., & K am, T. 1999.
Golden
eagle
p eda ion on semidomes ic eindee .
—
Wildli e
Socie y
Bulle in
27(4):
1038-1042.
Nybakk,
K.,
Kjel ik,
O., K am, T., O e skaug, K., &
Sunde,
P. 2002.
Mo ali y
o semi-domes ic eindee
Rangi e
a andus
in cen al
No way.
—
Wildli e
Biology
8(1): 63-68.
Ollila,
T. 2009. Rapo i maako kan, muu ohaukan,
un u ihaukan
seka
Oulun
ja
Lapin
laanien
me iko ki-
en
pesinnois a uonna 2009.
—
Me sdhalli us,
Luon o-
pal elu 2009,
8pp. (In
Finnish).
Rangi e ,
30
(1),
2010
87