DIURNAL DEFECATION RATE OF MOOSE IN SOUTHWEST FINLAND
Juho Ma ala
1
and An i Uo ila
2
1
Finnish Fo es Resea ch Ins i u e Me la, P.O. Box 68, FI-80101 Joensuu, Finland;
2
Uni e si y o Helsinki
Hyy iälä Fo es S a ion, Hyy iälän ie 124, FI-35500 Ko keakoski, Finland.
ABSTRACT: An accu a e measu e o de eca ion a e is essen ial o applica ion o pelle g oup coun s
in moose (Alces alces) popula ion es ima es. We measu ed he win e ime, diu nal de eca ion a e o
moose by acking 7 GPS-colla ed and 22 uncolla ed moose in sou hwes Finland. The mean de eca-
ion a e was 23.5 ± 4.2 pelle g oups/d, one o he highes alues epo ed. The mean de eca ion a e
did no di e be ween he acking me hods (GPS s. uncolla ed moose); limi ed sample size p e-
cluded conclusions abou sex and age di e ences. The de eca ion a e was no co ela ed wi h calenda
week, leng h o accumula ion pe iod, o numbe o diu nal beds. Ou esul s a e app op ia e o use in
sou hwes Finland when using he pelle g oup me hod o assess moose popula ion densi y.
ALCES VOL. 49: 155–161 (2013)
Key wo ds:Alces alces, Finland, de eca ion a e, moose, pelle g oup, acking.
Coun ing ecal pelle g oups o moose
(Alces alces) has been widely used o es i-
ma e habi a u iliza ion, eeding beha io ,
and popula ion ends and densi y (see
F anzmann e al. 1976b, Fo bes and The-
be ge 1993, Hä könen and Heikkilä 1999,
Rönnegå d e al. 2008, Månsson 2009, Mån-
sson e al. 2011a, b). Reliable moose popula-
ion es ima es a e no always ealized om
pelle g oup coun s (Rönnegå d e al.
2008), bu he me hod's use ulness has been
no ed (Ne 1968, Lau enschlage and Jo dan
1993, Månsson e al. 2011b), despi e some
unce ain y (Ne 1968). Howe e , o suc-
cess ully es ima e moose popula ion densi y
using a pelle g oup coun , i is c i ical o
use an accu a e de eca ion a e in he su ey
a ea and ime pe iod (Timme mann 1974).
Ea lie s udies ha e used 2 main me h-
ods o es ima e he de eca ion a es o
moose: 1) ack moose in snow-co e ed e -
ain (Joyal and Rica d 1986, Ande sen e al.
1992), and 2) es ima e he numbe o pelle
g oups in a closed a ea o island whe e he
numbe o moose is known (Jo dan e al.
1993). De eca ion a es ha e occasionally
been es ima ed by compa ing he ae ial cen-
suses and pelle g oup coun s in speci ic
a eas (Rönnegå d e al. 2008). Moose enclo-
su es could also be u ilized, bu esul s om
domes ic moose can be a ec ed by ood
quali y and beha iou ha a e dissimila o
na u al condi ions. GPS adio-colla s enable
in ensi e and accu a e acking by iden i y-
ing speci ic indi iduals and he beginning
and end poin s o hei speci ic acks, p o-
iding ideal condi ions o measu e de eca-
ion a es o ee- anging moose.
Moose de eca ion a es a y by age, sex,
habi a , ood quali y, season, and yea (Des-
Meules 1968, F anzmann e al. 1976a, Olde-
meye and F anzmann 1981, Joyal and
Rica d 1986, Ande sen e al. 1992, Månsson
e al. 2011b). La ge a ia ions in de eca ion
a e ha e been epo ed in di e en a eas;
o example, in No h Ame ica a ia ion
was 9.6–32.2 pelle g oups/d (Timme mann
1974), and in no he n Eu ope a es a ied
Co esponding au ho : Juho Ma ala, Finnish Fo es Resea ch Ins i u e Me la, P.O.Box 68, FI-80101
Joensuu, Finland. +358 40 801 5275. juho.ma a[email p o ec ed]
155
om 14–26.9 pelle g oups/d (Ande sen e al.
1992, Remm and Luud 2003, Rönnegå d
e al. 2008). These da a emphasize he
impo ance o using a ea-speci ic a es
when using he pelle g oup me hod o es i-
ma e moose popula ion densi y.
Ou main objec i e was o o mula e a
gene al es ima e o he win e ime, diu nal
de eca ion a e o moose in sou hwes Fin-
land by acking bo h GPS-colla ed and
uncolla ed moose in snowy e ain. We also
compa ed hese 2 acking me hods and
looked o di e ences in de eca ion a es
be ween sex and age.
STUDY AREA
Moose we e acked in 2 sepa a e a eas
app oxima ely 100 km apa in sou hwes
Finland (Fig. 1). Uncolla ed moose we e
acked in he O i esi-Kangasala a ea
(∼WGS84 61°36′N, 24°22′E) and GPS-
colla ed moose in he Loppi-Hy inkää a ea
(∼WGS84 60°38′N, 24°35′E). Bo h a eas
a e loca ed in he sou he n bo eal ege a ion
zone wi h Sco s pine (Pinus syl es is) and
No way sp uce (Picea abies) as he domi-
nan ee species. Fo es co e was 78% o
he o al land a ea in O i esi-Kangasala and
71% in Loppi-Hy inkää (Me la 2012).
METHODS
T acking o Uncolla ed Moose
We ac i ely sea ched o uncolla ed
moose in hei known habi a s du ing esh
snow condi ions be ween Decembe and
Ap il, 1999–2003. The accumula ion pe iod
o pelle coun s began when moose we e
seen o lushed, enabling da a collec o s o
accu a ely ime hei coun by loca ing esh
es ing places, pelle g oups, o acks in
Fig. 1. S udy a ea loca ions and s a ing poin s o moose acking pe iods in sou he n Finland. In he
O i esi-Kangasala a ea some o he s a ing poin coo dina es o uncolla ed moose a e ough
es ima es and o e lap because hey could no be sepa a ed a he map scale.
156
DIURNAL DEFECATION RATE OF MOOSE –MATALA AND UOTILA ALCES VOL. 49, 2013
he snow. The pelle g oups we e coun ed he
ollowing day, un il moose led om he
coun e . The end o he accumula ion pe iod
was de e mined in a simila manne as he
s a ing poin . The accumula ion pe iods
a ied om 8–31 h.
Moose we e classi ied as ei he adul s o
cal es by isually obse ing hem and hei
pelle s, bedding places, and beha io . Thei
sex could be de e mined wi hou isual con-
ac by analyzing hei u ina ion me hods,
as bulls u ina e in on o he hind hoo es.
We measu ed he diu nal de eca ion a e o
5 bulls, 3 cows, and 3 cal es. Addi ionally
we measu ed 3 cows wi h win cal es and
1 cow wi h a single cal , wi hou sepa a ing
he pelle g oups o he cows and cal es. In
o al, we we e able o eco d he diu nal de e-
ca ion a es o 22 uncolla ed moose and
coun he numbe o beds o 7 indi iduals.
T acking o GPS-Colla ed Moose
The Finnish Game and Fishe ies
Resea ch Ins i u e (FGFRI) p o ided loca-
ion da a o he GPS-colla ed moose. The
FGFRI implemen ed GPS-colla ing p oce-
du es in acco dance wi h Finnish legisla ion,
wi h pe mission om he Na ional Animal
Expe imen Boa d o Finland. An indi idual
GPS-colla ed cow was acked once and
ano he on 5 sepa a e occasions, while 1
cow wi h win cal es was acked once, and
one cow wi h a single cal wice; al oge he ,
7 indi iduals we e acked.
The GPS-colla ed moose we e acked a
ew days a e esh snow all in Decembe –
Ma ch 2010. The coun e wen o he mos
ecen loca ion o moose acks which we e
usually ∼12 h old. The pelle g oups we e
coun ed along he moose ack by ollowing
i agains he o iginal cou se o he moose;
he coo dina es o he pelle g oups, beds,
and u ina ion si es we e loca ed wi h hand-
held GPS de ices. The accumula ion pe iod
inished when an indi idual ack became
mixed wi h o he s o sunse p ecluded
acking. The du a ion o he accumula ion
pe iod was de e mined by iden i ying he
ime associa ed wi h he closes loca ion o
he acks wi h GPS colla da a. The accumu-
la ion pe iods anged om 6–47 h.
Da a Analysis
A leas 20 pelle s we e equi ed o make
a pelle g oup. We p ocessed he pelle g oup
da a o 29 moose (7 GPS-colla ed, 22 uncol-
la ed) o calcula e he diu nal de eca ion a e
(numbe o pelle g oups p oduced pe indi-
idual in 24 h) om he accumula ion pe i-
ods o indi idual bulls, cows, and cal es.
Mean alues we e calcula ed o he g oup
o cows and cal es when i was impossible
o iden i y cal om cow pelle g oups; ana-
lyses o 25 sepa a e cases we e used o calcu-
la e he diu nal de eca ion a e.
Fo compa ison, we sampled he mean
alues o diu nal de eca ion a es o 3 sex
and age classes: 1) bulls, 2) cows, and 3)
cal es and cow-cal es. The las class was
equi ed because we we e only able o ack
3 indi idual cal es, which was insu icien
o any easonable analysis. We also com-
pa ed he mean de eca ion a es o he uncol-
la ed and GPS-colla ed moose. Due o
limi a ions in he linea i y and homogenei y
o a iances in he da a, we used he
K uskal-Wallis es o mul iple samples
and he Mann-Whi ney es o pai ed sam-
ple compa isons. Fu he mo e, we sea ched
o possible co ela ions be ween he diu nal
de eca ion a e and 1) he calenda week o
he accumula ion pe iod, 2) he du a ion o
he accumula ion pe iod, and 3) he numbe
o diu nal beds o pa o he samples. All
s a is ical calcula ions we e pe o med using
he S a is ical Package o he Social
Sciences (SPSS) 17.0 so wa e.
Da a ela ing o home ange o moose
du ing he accumula ion pe iod (calcula ed
using he Home Range Tools o A cGIS®
e sion 1.1 wi h he Minimum Con ex
Polygon me hod), leng h o he moose ack
ALCES VOL. 49, 2013 MATALA AND UOTILA –DIURNAL DEFECATION RATE OF MOOSE
157
du ing he accumula ion pe iod, and he
numbe o diu nal u ina ions we e also col-
lec ed om he GPS-colla ed moose, bu
insu icien sample sizes p ecluded hei u i-
liza ion in he analysis.
RESULTS
The diu nal de eca ion a e anged om
12.2–32 pelle g oups wi h an o e all mean
o 23.5 ± 4.2 (SD; Table 1, Fig. 2). The
mean alues o bull, cow, and cal /cow-cal
g oups we e di e en (K uskal-Wallis es :
χ
2
= 9.9, d = 2, P= 0.007; Table 1). The
cal and cow-cal g oup had he highes
mean a e, bu was s a is ically di e en
only om he cow g oup (Table 1). The
bull g oup had he lowes mean a e, bu
also he lowes and highes absolu e alues
(i.e., he wides ange; Table 1). The cow
and bull g oup a es and he a es o he
uncolla ed and GPS-colla ed moose we e
no di e en (Table 1).
The diu nal de eca ion a e was no
ela ed o he calenda week o he accumula-
ion pe iod (Fig. 2, Table 2). The GPS-
colla ed indi idual which was acked 5
imes be ween 17 Janua y and 23 Feb ua y
2010 showed no ends du ing his pe iod.
The de eca ion a e did no co ela e o he
du a ion o he accumula ion pe iod o o
he numbe o diu nal beds (Table 2). The
mean alues o he o he a iables we e:
diu nal numbe o beds = 8.0 ± 4.2 (SD,
n = 15), he diu nal u ina ion a e = 1.0 ±
1.0 (SD, n = 9), he a ea o home ange du -
ing he accumula ion pe iod = 95,267 m
2
±
122,399 (SD, n = 9), and leng h o ack du -
ing he accumula ion pe iod = 955 m ± 832
(SD, n = 6).
DISCUSSION
The mean de eca ion a e was conside ed
high (23.5 ± 4.2 pelle g oups/d) bu simila o
he a e age a e measu ed in ela i ely good
moose habi a in sou he n No way (22.9 pel-
le g oups/d; Ande sen e al. 1992). Lowe
de eca ion a es we e measu ed in sou he n
Sweden (14 pelle g oups/d; Rönnebe g e al.
2008) and on Isle Royale, No h Ame ica
(20.9; Jo dan e al. 1993). Many s udies
ha e epo ed lowe de eca ion a es in
Alaska and Canada (see DesMeules 1968,
F anzmann e al. 1976a, Oldemeye and
F anzmann 1981, Joyal and Rica d 1986).
The high alues epo ed in ou s udy
a ea a e no un easonable when compa ing
he s a us o he moose popula ion in Finland
o o he No dic coun ies. The Finnish
moose popula ion is lowe and o a highe
p oduc i e s a e compa ed o hose in
Table 1. Mean alues o he win e ime, diu nal de eca ion a e and compa isons be ween moose g oups in
sou he n Finland.
Pelle g oups (#/ind/24 h) Mann-Whi ney Tes
Mean Min Max n SD Tes agains U- alue P
G ouping by moose ype:
Bulls 20.1 12.2 32.0 5 7.3 Cow 12 0.152
Cows 22.3 18.9 24.0 9 2.2 Cal + g oups 10 0.002
Cal and Cow-cal g oups 25.9 23.0 31.3 11 2.2 Bull 11 0.061
G ouping by acking me hod:
GPS-colla ed 23.5 20 26.7 9 1.8 Uncolla ed 64 0.647
Uncolla ed 23.4 12.2 32.0 16 5.2
All moose 23.5 12.2 32.0 25 4.2
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DIURNAL DEFECATION RATE OF MOOSE –MATALA AND UOTILA ALCES VOL. 49, 2013
Sweden and No way (La sund e al. 2003,
Tiilikainen e al. 2012). These di e ences
p esumably imply be e o aging habi a ,
highe nu i ional condi ion, and a esul an
highe de eca ion a e in Finland. Because
de eca ion a es a e in luenced by ood qual-
i y and a ailabili y and show la ge a ia ion
(Ande sen e al. 1992), i ollows ha a ea-
speci ic de eca ion a es should be measu ed
when using pelle g oup coun s o es ima e
popula ion densi y.
The ex emes in de eca ion a e a ied
la gely in ou s udy, bu mos obse a ions,
especially o he GPS-colla ed indi iduals,
we e simila o he mean indica ing he gen-
e al eliabili y o hese da a and hei applica-
ion o es ima e popula ion densi y. The
ex eme alues could esul om local o a-
ging condi ions o om longe mo emen s
associa ed wi h unin ended dis u bances
while acking he uncolla ed moose.
Because he de eca ion a e can co ela e
wi h age and sex, hese ela ionships should
be aken in o accoun i a change in popula-
ion s uc u e occu s (F anzmann e al.
1976a). Ou da a indica e ha he cal and
cow-cal g oup had sligh ly highe de eca-
ion a es han indi idual cows as also
epo ed by DesMeules (1968), bu opposi e
ha o Joyal and Rica d (1986). Bulls and
cows we e no di e en in ou analyses;
howe e , gene al conclusions abou sex and
age di e ences canno be made due o he
limi ed sample size.
Using an accu a e de eca ion a e is c i i-
cal when applying pelle g oup coun s in
moose popula ion densi y es ima es. De eca-
ion a es ha e no been published p e iously
in Finland, and gi en he conside able a ia-
ion in simila da a om no he n Eu ope, we
conside ou esul s he bes a ailable o
local use. Fu he mo e, simila de eca ion
a es we e ob ained o bo h egions in ou
s udy, making he mean alues applicable
o all o sou he n Finland. The lack o co e-
la ion be ween calenda week and de eca ion
a e indica es a s able accumula ion a e
h oughou win e , sugges ing ha hese
a es migh be applicable o sp ing ime pel-
le g oup coun s. Va ia ions in he absolu e
da a and means should be aken in o accoun
when calcula ing con idence limi s o he
inal moose densi y es ima e.
ACKNOWLEDGEMENTS
We hank M . J. Sillanpää o acking
he GPS-colla ed moose and M . H. Ma ila
50
Fig. 2. The diu nal de eca ion a e o moose in
sou he n Finland ela i e o he ime o he
acking pe iod (calenda weeks). Moose ypes
a e classified as: 1) indi idual bulls, cal es,
and cows, and 2) cow-cal g oups (when
indi iduals could no be sepa a ed du ing
acking); cow1 = cow wi h single cal ,
cow2 = cow wi h 2 cal es.
Table 2. Pea son co ela ions be ween he diu nal
de eca ion a e (pelle g oups/moose/24 h) and
empo al a iables in sou he n Finland; no
di e ences we e ound.
Calenda
week
Accumula ion
pe iod (h)
Beds/
moose/
24 h
Coe icien −0.144 −0.086 −0.403
P- alue 0.492 0.682 0.136
n25 25 15
ALCES VOL. 49, 2013 MATALA AND UOTILA –DIURNAL DEFECATION RATE OF MOOSE
159

o acking in he Kangasala a ea. D . J.
Pusenius om he Finnish Game and Fish-
e ies Resea ch Ins i u e is g ea ly acknowl-
edged o o ganizing he GPS-colla ing and
hus enabling da a a ailabili y on exac
moose loca ions o ou s udy. We also hank
he Finnish Fo es Resea ch Ins i u e, he
Uni e si y o Helsinki, and he Minis y o
Ag icul u e and Fo es y o p o iding inan-
cing o ou wo k. M.Sc. M. Melin is
acknowledged o p oducing he maps in
Fig. 1 and Ms. S.Thompson o checking
he language o his manusc ip .
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