RESEARCH ARTICLE Open Access
Applica ion o change-poin analysis o de e mine
win e sleep pa e ns o he accoon dog
(Nyc e eu es p ocyonoides) om body empe a u e
eco dings and a mul i- ace ed die a y and
beha io al s udy o win e ing
Anne-Ma i Mus onen
1,2*
, Te u Lempiäinen
3
, Mikko Aspelund
4
, Paa o Hells ed
5
, Ka i Ikonen
2
, Juhani I ämies
6
,
Ville Vähä
7
, Jaakko E kina o
7
, Juha Asikainen
2
, Me i Kunnas an a
8
, Pekka Niemelä
9
, Ja i Aho
10
and
Pe e i Nieminen
1,2
Abs ac
Backg ound: A mul i- ace ed app oach was used o in es iga e he win e ime ecophysiology and beha io al
pa e ns o he accoon dog, Nyc e eu es p ocyonoides, a sui able model o win e sleep s udies. By u ilizing GPS
acking, ac i i y senso s, body empe a u e (T
b
) eco dings, change-poin analysis (CPA), home ange, habi a and
die a y analyses, as well as a y acid signa u es (FAS), he impac o he species on win e ime ood webs was
assessed. The iming o passi e bou s was de e mined wi h mul iple me hods and compa ed o T
b
da a analyzed
by CPA.
Resul s: Raccoon dogs displayed win e ime mobili y, and he home ange sizes de e mined by GPS we e simila
o la ge han p e ious es ima es by adio acking. The p e e ed habi a s we e ga dens, sho es, deciduous o es s,
and spa sely o es ed a eas. Fields had close o neu al p e e ence; oads and ail oads we e u ilized as a el ou es.
Raccoon dogs pa icipa ed ac i ely in he ood web and gained bene i om human ac i i y. Mammals, plan s,
bi ds, and disca ded ish comp ised he mos impo an die a y classes, and he consump ion o ish could be
de ec ed in FAS. Ambien empe a u e was an impo an ex e nal ac o in luencing T
b
and ac i i y. The iming o
passi e pe iods app oxima ed by beha io al da a and by CPA sha ed 91% simila i y.
Conclusions: Passi e pe iods can be de e mined wi h CPA om T
b
eco dings wi hou he p e iously used
ime-consuming and expensi e me hods. I would be possible o ec ui mo e animals by using he simple
me hods o da a logge s and ea ags. Hun ing could be used as a ool o e u n he ea - agged indi iduals
allowing he economical ex ension o ollow-up s udies. The T
b
and CPA me hods could be applied o o he
no he n ca ni o es.
Keywo ds: Body empe a u e, Change-poin analysis, Fa y acid signa u e, Fo aging ecology, GPS acking, Home
ange, Nyc e eu es p ocyonoides, Win e sleep
* Co espondence: [email p o ec ed]
1
Ins i u e o Biomedicine/Ana omy, School o Medicine, Facul y o Heal h
Sciences, Uni e si y o Eas e n Finland, P.O. Box 1627, FI-70211, Kuopio,
Finland
2
Depa men o Biology, Facul y o Science and Fo es y, Uni e si y o Eas e n
Finland, P.O. Box 111, FI-80101, Joensuu, Finland
Full lis o au ho in o ma ion is a ailable a he end o he a icle
© 2012 Mus onen e al.; licensee BioMed Cen al L d. This is an Open Access a icle dis ibu ed unde he e ms o he C ea i e
Commons A ibu ion License (h p://c ea i ecommons.o g/licenses/by/2.0), which pe mi s un es ic ed 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.
Mus onen e al. BMC Ecology 2012, 12:27
h p://www.biomedcen al.com/1472-6785/12/27
Backg ound
To yield eliable da a no in luenced by human dis u -
bance, se e al me hods ha e been de eloped o moni o
ac i i y pa e ns o wild mammals. Ve y high equency
(VHF) acking is mos o en u ilized bu i is ime-
consuming and labou -in ensi e. This makes he use o
adio eleme y expensi e, al hough adio colla s a e gene-
ally qui e cheap and hei longe i y is good. Global posi-
ioning sys em (GPS) acking yields mo e abundan and
accu a e home ange da a bu he equipmen is expensi e
and ails mo e easily [1]. Al hough GPS sys ems cos mo e
han VHF colla s, he cos pe one loca ion is o en smal-
le , as pe sonnel cos s o ield wo k can be minimized.
Moni o ing win e ac i i y is essen ial o desc ibing
he beha io o species u ilizing a complex win e ing
s a egy consis ing o al e na ing pe iods o physical ac-
i i y and passi i y. De e mining he du a ion and iming
o passi e bou s can be use ul in se e al ways. Win e
sleep pa e ns could be used as an indica o o clima e
change scena ios o moni o he e ec s o global wa m-
ing on bo eal ecosys ems. The clima e change can lead
o highe o aging e o in win e inc easing in a- and
in e species in e ac ions and, hus, also he ansmission
isk o abies, o he zoonoses, and pa asi es [2]. Fo his
eason, ac i i y o passi ely win e ing species should be
e-e alua ed wi h mode n acking me hods.
A e y use ul model species is he palea c ic accoon
dog (Nyc e eu es p ocyonoides), an in asi e omni o e,
which inhabi s empe a e o suba c ic egions. I is e y
abundan , ha es ed legally, and due o i s ela i ely small
body size, i can be handled sa ely and, e.g., anes hesia is
no needed o p ocedu es such as blood sampling. In
Finland, he accoon dog is conside ed an alien spe-
cies and i s popula ion size has showed an inc easing
end [3]. Recen ly, i s a ea o dis ibu ion has expanded
qui e apidly o he no he nmos Lapland, and Finnish
indi iduals a e also colonizing Sweden ia he no he n
ou e. The goals o he popula ion con ol include he p e-
en ion o he sp ead and es ablishmen o he species o
o he pa s o Scandina ia, as i is a ec o o diseases and
pa asi es and a po en ial h ea o na i e auna. I has also
been ponde ed i he sp ead o he dis ibu ion a ea o he
accoon dog could bene i om global wa ming [4]. The
ecophysiology o o e win e ing o he species was p e i-
ously in es iga ed wi h adi ional me hods [5,6]. As he
accoon dog displays an in e media e win e ing s a egy
be ween bea s (U sus spp.) and ac i ely win e ing ca ni-
o es (e.g., he ed ox Vulpes ulpes), s udying i s ecophy-
siology can inc ease he knowledge on he e olu ion o
di e en ypes o win e ing.
Fo hese easons, i would be desi able o be able o
de ine he es ing pe iods o animals economically wi h-
ou comp omising he eliabili y o he da a. The iming
o win e sleep bou s was p e iously app oxima ed om
he co e body empe a u e (T
b
) da a o accoon dogs [5].
Due o his, i would be possible o equip animals wi h
inexpensi e in a-abdominal empe a u e logge s and
ea ags, and o iden i y he pe iods o passi e win e ing
om he T
b
da a o each indi idual. To be able o pe -
o m a mo e de ailed analysis o he pa e ns o al e na -
ing passi i y and ac i e o aging bou s, a eliable
ma hema ical applica ion ha would be able o de ec
mo e sho - e m and sub le changes in T
b
hy hms and
ha would be less dependen on subjec i e human as-
sessmen could be mos use ul.
Theaimo hep esen se o expe imen swas oconduc
a comp ehensi e s udy on se e al p e iously neglec ed
aspec s o o e win e ing o he accoon dog. The species
could ha e a signi ican impac on ood webs du ing
i s ac i i y bou s in win e . This can be assessed i)by
de e mining he u iliza ion o di e en bio opes and
oad/ ail oad ne wo ks by posi ioning da a and habi a
use analysis. The win e ime die o he species can
be in es iga ed ii) by analyzing he con en s o gas oin es-
inal ac s oge he wi h adipose issue a y acids (FA)
and FA signa u es (FAS). iii) Ex e nal ac o s ha change
du ing global wa ming and a ec he iming and du a ion
o passi i y can also be de e mined. Finally, i )ama he-
ma ical me hod analyzing T
b
pa e ns can de e mine he
ime pe iods when accoon dogs ope a e ac i ely in win e -
ime ood webs. The hypo heses o he s udy we e as ol-
lows: i) abio ic ac o s in win e (ambien empe a u e T
a
,
snow) can a ec he habi a use o accoon dogs and di ec
hem o u ilize oad/ ail oad ne wo ks as pa hways o o -
aging, ii) accoon dogs u ilize ood esou ces oppo unis-
ically du ing win e ime ac i i y bou s and can ha e an
impac on ood webs in hei habi a s, iii) ac o s liable o
he clima e change (T
a
, snow dep h) can ha e an e ec on
he ac i i y pa e ns and, hus, win e sleep could be
suscep ible o global wa ming, and i )T
b
pa e ns
can be subjec ed o ma hema ical analysis in e p e ing
eliably he passi e pe iods o win e ing om he T
b
da a o wild accoon dogs.
The p esen s udy de eloped a p ac ical ool o assess
om easily ob ainable da a (in a-abdominal T
b
) he pe -
iods o passi i y, he de e mina ion o which would
o he wise equi e ei he ex ensi e wo k in he ield
( adio acking) o expensi e equipmen (GPS colla s
and ac i i y senso s combined wi h da a ans e cos s).
This no el and economical me hod u ilizing change-
poin analysis (CPA) can be used no only in esea ch
p ojec s on he accoon dog bu e en ually applied also
o o he passi ely win e ing ca ni o es.
Me hods
Body empe a u e, ac i i y, and home anges
The s udy p ocedu es we e app o ed by he Animal
Ca e and Use Commi ee o he Uni e si y o Joensuu in
Mus onen e al. BMC Ecology 2012, 12:27 Page 2 o 17
h p://www.biomedcen al.com/1472-6785/12/27
2006–2007 and by he Finnish Na ional Animal Expe i-
men Boa d in 2008–2010 (#ESLH-2008-06316/Ym-23),
and complied wi h he cu en laws o Finland. Fou een
accoon dogs we e cap u ed li e wi h box aps, dogs, o
using a cable in Kaa amo and Ris inkylä illages in
eas e n Finland (62.56116338 N; 29.14269194 E) in au-
umn 2006 and 2007. They we e anes he ized wi h in a-
muscula ke amine (5 mg/kg) and xylazine (2 mg/kg) and
wo s e ile he mosensi i e da a logge s (iBu on The mo-
ch on DS1921H, Maxim In eg a ed P oduc s, Sunny ale,
CA) egis e ing he T
b
a 120-min in e als we e implan ed
in o hei abdominal ca i ies [5]. The accu acy and p eci-
sion o he logge s had been es ed igo ously [7]. The ani-
mals we e i ed wi h ea ags (model 1841, Na ional Band
& Tag Co, Newpo , KY) and s o e-on-boa d GPS colla s
wi h emo e GSM and a back-up VHF adio beacon
(Tellus Basic colla 2A, Followi AB, S ockholm, Sweden;
weigh 3–4% o body mass, BM). The GPS colla s we e
p og ammed o eco d 8 posi ion coo dina es pe day (a
01:00, 02:00, 04:00, 10:00, 20:00, 22:00, 23:00, 24:00 h)
based on he mos ly noc u nal ac i i y pa e n o he spe-
cies [2]. One day ime ix was ob ained o posi ion he es
si e. The ac i i y senso s measu ed he change in he acce-
le a ion o he colla in wo axes (x, y) du ing each 120-s
ime pe iod used o ob ain a GPS ix. No exac anges o
ac i i y alues indica ing ac i i y s. passi i y we e de e -
mined p e iously o he s udied species, and o he
p esen expe imen , he ac i i y sco e sum [Σ(x + y)] was
calcula ed o each da e o each indi idual.
BM and body leng hs we e de e mined and body mass
indices (BMI) co ela ing wi h he body a -% [8] calcu-
la ed. Age was es ima ed by palpa ion o he p ominence
o he ulna (closu e o he epiphyseal pla e) and, i a hun ed
animal was deli e ed o he Uni e si y, pos mo em wi h
his ological examina ion o a canine oo h oo [9]. The
indi iduals <1 yea o age we e conside ed ju eniles and
hose >1 yea o age we e classi ied as adul s. A blood sam-
ple was aken om a saphenous ein o a hind leg wi h
s e ile needles and sy inges wi h e hylenediamine e aace-
ic acid, and he comple e blood coun (CBC) was de e -
mined as an indica o o heal h wi h he Ve abc Animal
Blood Coun e (ABX Hema ologie, Mon pellie , F ance),
adjus ed o he canine hema ologic p o ile. A sample
(1–2 g) o en al subcu aneous (sc) a was emo ed
and s o ed a –80°C. A e a eco e y pe iod o app oxi-
ma ely one week a a u a m, he animals we e eleased
a he cap u e si es, ecap u ed li e o dead in he ollow-
ing sp ing, and he li e indi iduals we e ope a ed simila ly.
Mo eo e , 6 accoon dogs we e cap u ed, ope a ed, and
eleased uncolla ed in au umn 2007, 2008, and 2009. They
we e e u ned dead by hun e s in he subsequen sp ings,
and he da a logge s we e eco e ed. Taken oge he , T
b
logge s we e e ie ed om 10 colla ed animals [2 adul
males (M1, M3), 3 ju enile males (M2, M5, M6), 3 adul
emales (F3, F4, F6), 2 ju enile emales (F1, F2)] and 6
uncolla ed animals [1 adul male (M9), 1 ju enile male
(M8), 1 adul emale (F11), 3 ju enile emales (F8, F9,
F10)]. GPS colla s we e eco e ed om 12 indi iduals
[3 adul males (M1, M3, M4), 3 ju enile males (M2, M5,
M6), 3 adul emales (F3, F4, F6), 3 ju enile emales (F1,
F2, F5)], as wo colla s go dys unc ional.
All successi e eloca ions we e included in he home
ange analyses al hough hey may ha e been au oco e-
la ed, i.e., aken oo closely in ime o be s a is ically inde-
penden , o ge he bes a ailable es ima es [10-12]. The
ixed Ke nel me hod (K95/50%; [13]) was used o es ima e
he home ange sizes and co e a eas (50% o he loca ions)
oge he wi h he inc emen al analysis using he Ranges7
so wa e (Ana ack L d, Wa eham, UK; [14]). Two ju e-
niles pe o med long-dis ance dispe sion om one home
ange o ano he . Fo F2, he eloca ions o hese home
anges we e analyzed sepa a ely, and o M5, only he pos -
dispe sal eloca ions we e included in he home ange ana-
lysis as, based on he inc emen al analysis and p e ious
da a [15], he numbe o ixes in he ini ial home ange
was insu icien . The calcula ed o al home ange a eas
we e plo ed on a habi a map (CLC2000, Eu opean En i -
onmen Agency, Copenhagen, Denma k) in a geog aphic
in o ma ion sys em (A cGIS 10, Es i, Redlands, CA) o as-
sess he ela i e impo ance o each habi a ype du ing
win e . Habi a selec ion was analyzed by de e mining,
i s ly, which habi a s we e used based on he eloca ions
o he indi iduals. Secondly, his was compa ed o he
o e all habi a dis ibu ion in he home ange. A selec ion
index [(% o eloca ions)/(% o po en ially a ailable habi-
a )] was calcula ed o each habi a ype o de e mine,
which habi a s we e a o ed o anked low.
The win e ime T
b
eco dings (p incipally in No –
Ma ch) we e di ided in o ac i e and passi e pe iods based
on he abili y o he colla o posi ion he animal, eloca-
ions, and ac i i y sco es. The (x + y)- alues o he ac i i y
senso s we e e alua ed oge he wi h he posi ioning da a
o assess ac i i y/passi i y (ac i i y sco es + posi ioning =
beha io al da a). An animal was conside ed passi e when i
s ayed a he same GPS posi ion, i s colla could no es ab-
lish sa elli e connec ion (i.e., he animal was mos likely
below he g ound o hiding in dense unde g ow h ege a-
ion), and/o when i s 24-h ac i i y sco es we e close o
ze o o , based on he gene al appea ance o indi idual
ac i i y da a, lowe han du ing he nigh s wi h docu-
men ed o aging. Snow dep h was moni o ed once a week
and a e each snow all, and he mosensi i e logge s
(iBu on The moch on DS1921G) synch onized wi h he
T
b
eco dings egis e ed he T
a
a 2-h in e als.
Die a y analyses and a y acid signa u es
The composi ion o he die was s udied om 93 esh
ca casses ha es ed du ing he pe iods o snow co e
Mus onen e al. BMC Ecology 2012, 12:27 Page 3 o 17
h p://www.biomedcen al.com/1472-6785/12/27
be ween No 5 2006–Ap il 13 2007 and No 9 2007–
Ma ch 30 2008. Ca casses we e sexed and weighed, and
he hickness o he en al sc a and he mass o he
omen um we e measu ed. The gas oin es inal ac was
dissec ed, weighed, and ozen a –20°C. I he animal
had been killed <24 h s ea lie and s o ed ozen, 1–2g
o en al sc a was dissec ed and s o ed a –80°C o
FAS analysis.
A e hawing, he s omachs and in es ines we e sepa-
a ed, weighed ull, opened, insed in a sie e (mesh size
0.5 mm), and weighed emp y. Thei con en s we e
s o ed in E OH. Expe s iden i ied he undiges ed ood
emains ( ee h, bones, hai s, ea he s, seeds, e c.) mac o-
and mic oscopically o he lowes possible axon [16-19].
No mic oscopic sea ch was pe o med o ea hwo m
che ae as g ound os p esumably made he access o
ea hwo ms e y unlikely, and hei impo ance was p e-
iously negligible in win e die [20]. Hai samples we e
laid on a s ip o balsa wood and ixed using colou less
nail a nish. D ied samples we e cu wi h a azo blade,
he c oss-sec ion was iewed mic oscopically, and he
species/genus iden i ied [21,22]. The olumes o ood
emains we e measu ed using glass measu ing cylinde s.
The ood i ems we e classi ied in o 13 ca ego ies: ce eals,
be ies, ege ables, ui s, oden s, insec i o es, ca ni o es,
lepo ids, ce ids, bi ds, ish, in e eb a es, and o he di-
ges ible ma e ial ha could no be u he classi ied. The
o al o use ul plan s was calcula ed as he sum o ag icul-
u al and u ili y plan s. Likewise, sums o wild, diges ible,
and undiges ible plan s, and small, medium-sized, and
la ge mammals we e calcula ed.
The esul s a e p esen ed as he olume (ml) o each
ood i em in he s omachs/in es ines and as he olume
o each ood i em o he o al olume o he s omach/
in es inal ood i ems ( ela i e sha e, RS%). Man-made
ma e ial, undiges ible plan ma e ial, soil, and small
amoun s o accoon dog hai s p esumably inges ed du ing
g ooming we e excluded, as we e he inges ed bai s om
he aps. Vol1 s ands o he o al olume o all diges ible
ood i ems excluding bai s, whe eas Vol2 signi ies he o al
olume o all inges ed ma e ial including nondiges ible
cons i uen s, bai s, and g oomed accoon dog hai . The
equency o occu ence (FO) was calcula ed as ollows:
i) FO1= 100 × he p opo ion o s omachs/in es ines con-
aining each ood i em and ii)FO2=100× heoccu ence
o each ood i em/ he o al numbe o occu ences o all
ood i ems. The die di e si y (numbe o di e en ood
i ems pe s omach/in es ine) and he main ood ( he mos
oluminous ood ype pe s omach/in es ine) we e also
de e mined. Diges i e ac con en s we e examined
mac oscopically o oundwo ms.
The FA composi ion e lec ing he mo e long- e m
die a y habi s was de e mined om he sc a o T
b
logge -
implan ed accoon dogs in 2004–2008 (n = 52; [5,23]) and
o eshly ozen ca casses in 2006–2008 (n = 33). The sam-
ples we e ansme hyla ed by hea ing wi h 1% H
2
SO
4
in
me hanol unde ni ogen a mosphe e, he FA me hyl es e s
we e ex ac ed wi h hexane and analyzed by a gas–liquid
ch oma og aph (6890N; Agilen Technologies Inc, San a
Cla a, CA) as desc ibed p e iously [23]. Rela i e changes in
he p opo ions o FA du ing win e ing we e calcula ed
by he o mula [(mol-% in sp ing)–(mol-% in au umn)]/
[mol-% in au umn]. These we e calcula ed o he indi-
iduals ha could be cap u ed bo h in au umn and
subsequen sp ing (n = 14), while he seasonal FA p o-
iles we e de e mined om all animals (au umn n = 39,
sp ing n = 46).
Change-poin analysis o body empe a u es
The ime pe iods o passi i y de e mined by beha io al
da a we e compa ed o passi e pe iods de ined om T
b
da a by CPA. I is a powe ul ool o de ec ing changes in
a ime-se ies. A simple p ocedu e o pe o ming CPA
was in oduced by Taylo [24] based on he assump ion o
he mean-shi da a. X
1
,X
2
,... ep esen he da a in ime-
o de . The mean-shi model can be w i en as X
i
=μ
i
+ε
i
,
whe e μ
i
is he a e age a ime iand ε
i
is he andom e o
associa ed wi h he i
h
alue. Gene ally μ
i
=μ
i−1
excep o
a small numbe o alues o icalled he change-poin s. I
is assumed ha andom e o e ms ε
i
a e independen
and hei means a e ze o. The pa e n es was u ilized o
ecognizing he mean-shi da a [25].
The p ocedu e o pe o ming CPA uses a combi-
na ion o cumula i e sum (CUSUM) cha s and boo -
s apping o de ec he changes. CUSUM cha s a e
cons uc ed by calcula ing a CUSUM based on he da a.
These sums a e he CUSUM o he di e ences be ween
he alues and he a e age. When X
1
,X
2
,...,X
n
ep e-
sen he da a, he CUSUM a e calcula ed as ollows: 1)
calcula ing he a e age o he da a 
X, 2) s a ing he
CUSUM a ze o by se ing S
0
= 0, and 3) calcula ing he
o he CUSUM by adding he di e ence be ween he
cu en alue and he a e age o he p e ious sum: S ¼
S 1þX 
XðÞ o =1,...,n. These CUSUM alues
p oduce a ime-se ies S
0
,...,S
n
. The g aph o his se ies
(CUSUM cha ) can be used o in e p e ing he occu -
ence o one o mo e changes. A segmen o he
CUSUM cha wi h an upwa d slope indica es a pe iod
whe e he alues end o be abo e he o e all a e age
and ice e sa. A sudden change in he di ec ion o he
CUSUM se ies indica es a sudden shi in he a e age.
A con idence le el o a possible change can be pe -
o med by a boo s ap analysis, o which an es ima o
o he magni ude o he change is equi ed. We used S
di
,
de ined as S
di
=S
max
−S
min
,whe eS
max
is he maximum
and S
min
he minimum o he CUSUM se ies. A single
boo s ap analysis can be pe o med by 1) gene a ing a
boo s ap sample, deno ed X
1
0
,...X
n
0
,by andomly
Mus onen e al. BMC Ecology 2012, 12:27 Page 4 o 17
h p://www.biomedcen al.com/1472-6785/12/27
eo de ing he o iginal alues X
1
,...,X
n
, 2) calcula ing he
boo s ap CUSUM based on he boo s ap sample,
deno ed S
0
0
,...,S
n
0
, 3) calcula ing he S
di
0
, and 4) de e mi-
ning, whe he he boo s ap di e ence S
di
0
is less han he
o iginal di e ence S
di
.
The boo s ap samples ep esen he case whe e
changes ha e no occu ed. By pe o ming a la ge num-
be o boo s ap samples and compa ing he es ima o s
S
di
0
wi h S
di
, i can be de e mined, a which p obabili y
a leas one change has occu ed. The con idence le el is
calcula ed as 100 × X/N%, whe e Nis he numbe o
boo s ap samples and Xis he numbe o boo s aps o
which S
di
0
<S
di
. A 95% con idence le el is equi ed o
de e mining ha a change has occu ed. Once a change
has been de ec ed, he iming o he change can be es i-
ma ed wi h he mean squa e e o (MSE) es ima o . The
MSE(m) is de ined as:
MSE mðÞ¼
X
m
i¼1
Xi
X1
ðÞ
2þX
n
i¼mþ1
Xi
X2
ðÞ
2
;
whe e 
X1¼
X
m
i¼1
Xi
mand 
X2¼
X
n
i¼mþ1
Xi
nm:
The MSE es ima o is based on he spli ing o he
da a in o wo segmen s, 1 o mand m+1 o n, es ima-
ing he a e age o each segmen and checking how well
he da a i he wo es ima ed a e ages. The alue o m
ha minimizes he MSE(m) is he bes es ima o o he
las poin be o e he change, and m+ 1 es ima es he
i s poin a e he change. Once a change has been
de ec ed, he da a can be di ided in o wo segmen s, one
on each side o he change-poin . Then he analysis is
epea ed o each segmen . Fo each signi ican change
ound, he segmen s con inue o be spli in wo, e c., o
be able o de ec mul iple changes.
CPA was applied o he T
b
da a o de ec ing he pe i-
ods o passi i y by sepa a ing he da a in o dis inc seg-
men s. The o iginal da a did no necessa ily sa is y he
mean-shi assump ion as consecu i e alues may ha e
been co ela ed. This co ela ion could be elimina ed by
handling he se ies o a e ages o wo consecu i e alues.
The ans o med da a s ill included a pe iodic s uc u e
caused by he 24-h hy hm o T
b
, and o his eason,
he e was a con lic wi h he assump ion o independen
e o s uc u e. To ci cum en his p oblem, he da a
we e sepa a ed in o sho o e lapping segmen s, as when
he segmen s a e sho enough, he e is no s ong e i-
dence o pe iodici y in each segmen . By applying CPA
o hese segmen s, he whole o iginal da a could be
di ided in o segmen s wi h di e en a e ages. The pe i-
ods wi h he lowes a e ages and he pe iods wi h ela-
i ely low a e ages su ounded by pe iods wi h highe
a e ages we e mos p obably he passi e pe iods. The
CPA applica ion was de eloped by using he MATLAB
p og am ( R2008a, Ma hWo ks, Na ick, MA). An ex-
ample o he ou pu can be examined in Figu e 1 ep e-
sen ing he o iginal T
b
da a o M9.
S a is ical analyses
Di e ences in he gene al a iables be ween he mon hs
we e analyzed wi h he one-way analysis o a iance
(ANOVA) o he nonpa ame ic K uskal–Wallis ANOVA
(SPSS 16.0 so wa e package, SPSS Inc, Chicago, IL). The
homogenei y o a iances and no mali y o dis ibu ion
we e es ed wi h he Le ene’s and Kolmogo o –Smi no
es s, espec i ely. The ac i i y sco e sum and a e age
24-h T
b
we e calcula ed o each da e o each accoon
dog, and indi idual ampli ude spec a we e calcula ed
wi h he Fas Fou ie T ans o m. The alues du ing
ac i e and passi e win e ing we e compa ed wi h he
independen samples S uden ’s - es o Mann–Whi ney
U- es o pa ame ic and nonpa ame ic da a, espec -
i ely. The la e was used when no mali y o dis ibu ion
was no a ained. The occu ence o ood i ems in he
gas oin es inal ac s was es ed wi h he χ
2
- es . To
analyze he empo al changes in he die , win e was
di ided in o No –Dec ( a s o age comple ed and pe iods
o passi i y begin o occu g adually), Jan–Feb (low o -
aging ac i i y), and Ma ch–Ap il (inc easing ac i i y and
ma ing season). To analyze he in e ela ionships be ween
he a e age 24-h T
b
and co a ia es T
a
, day leng h, and
snow dep h, he analysis o co a iance (ANCOVA) was
pe o med wi h he linea mixed model analysis. The
model included he indi idual as a andom ac o and he
co a ia es T
a
, day leng h, snow dep h, and bea ing cap-
aci y as well as hei in e ac ions wi h he indi idual. To
analyze he ela ionships be ween he FAS and di e en
die a y classes, he da a we e subjec ed o he mul i a ia e
p incipal componen analysis (PCA) using he SIRIUS
6.5 so wa e package (Pa e n Recogni ion Sys ems AS,
Be gen, No way; [26]). Bi a ia e co ela ions we e calcu-
la ed wi h he Spea man co ela ion coe icien (
s
). P<
0.05 was conside ed s a is ically signi ican . The esul s
a e p esen ed as he mean ± SE.
Resul s
Body empe a u e pa e ns and hei ma hema ical
analysis
Acco ding o CPA, he a e age numbe o passi e pe i-
ods was 7 ± 0.6, he sum o passi e days was 50 ± 5.4,
and he du a ions o he sho es and longes passi e
pe iods we e 1 ± 0.02 and 31 ± 7.1 days, espec i ely, in
he GPS colla ed indi iduals acked o 3–5 mon hs in
No –Ap il (G oup 1, n = 7). In hese animals, he com-
pa ibili y o he iming o he passi e pe iods, de e mined
by CPA and by beha io al da a, was 91% (78–100%). The
Mus onen e al. BMC Ecology 2012, 12:27 Page 5 o 17
h p://www.biomedcen al.com/1472-6785/12/27

a e age numbe o passi e pe iods (8 ± 1.2), he sum o
passi e days (56 ± 4.2), and he du a ions o he sho es
and longes passi e pe iods (3 ± 0.7 and 21 ± 4.2 days) o
he uncolla ed animals (G oup 2, n = 5) T
b
- eco ded bu
no acked du ing he same ime pe iod did no di e
om G oup 1, excep o he du a ion o he sho es
passi e pe iod (K uskal–Wallis ANOVA, H=9.467,d =2,
p< 0.01). The animals ha could be acked o only
sho e pe iods o ime (1–3 mon hs) du ing No –Feb
(killed by hun e s, dogs, o a ic; G oup 3, n = 4) dis-
played, on a e age, 1 ± 0.5 passi e pe iods, he o al leng h
o passi e win e ing was 6 ± 2.4 days, and he sho es
and longes pe iods las ed o 3 ± 0.9 and 7 ± 1.5 days
(K uskal–Wallis ANOVA, H=6.307–9.467, d = 2, p<
0.01–0.05 s. G oup 1). The compa ibili y o he iming
o he passi e pe iods de e mined by CPA and by beha-
io al da a a ied sligh ly mo e han in G oup 1 (mean
83%, ange 63–100%).
Gene ally, he longes passi e pe iod occu ed in
Jan–Ma ch. I was p eceded by 1–6 (3.8 ± 0.6) and
ollowed by 0–8 (2.6 ± 0.7) sho e passi e pe iods.
The o al du a ion o passi i y (days) co ela ed wi h
he numbe o passi e pe iods (
s
= 0.620, n = 16, p<
0.01) and wi h he du a ion o he longes passi e
pe iod (
s
= 0.810, n = 15, p< 0.001). Acco ding o he
ac i e/passi e classi ica ion by CPA, he a e age T
b
was 37.4 ± 0.08°C du ing ac i e and 36.5 ± 0.08°C du -
ing passi e win e ing, he di e ence being 1.0 ± 0.07°C
(Mann–Whi ney U- es , U= 4.000, n = 15, p< 0.001).
The e we e no di e ences in he a e ages be ween
he s udy g oups. The spec al analysis e ealed clea
24-, 12-, and 8-h oscilla ions in he T
b
o all animals
wi hou signi ican di e ences in he magni ude be-
ween he ac i e and passi e pe iods (Figu e 2). The
a e age 24-h T
b
was 37.6 ± 0.03°C in No –Dec, 36.7 ±
0.03°C in Jan–Feb, and 37.3 ± 0.07°C in Ma ch–Ap il,
all signi ican ly di e en om each o he (K uskal–
Wallis ANOVA, H= 87.716, d = 2, p< 0.001). The
a e age 24-h T
b
showed posi i e co a iance wi h he
a e age 24-h T
a
(ANCOVA, F
1,8.001
= 19.489, p<0.01),
while he in e ac ions wi h he o al dep h o snow,
dep h o so snow, and day leng h we e nonsigni i-
can (Figu e 3). The a e age 24-h T
b
o he wo indi-
iduals win e ing oge he (F8, M8) co ela ed
posi i ely (
s
= 0.854, n = 87, p< 0.001), and he iming
o hei passi e pe iods de e mined by CPA was 89%
iden ical.
Ac i i y pa e ns
The a e age ac i i y sco e was 86% lowe du ing pas-
si e o e win e ing han du ing he ac i e pe iods
de ined by CPA ( - es , | | = 14.259, n = 64, 80, p<
0.001). Du ing he ac i e pe iods o win e ing, he
a e age ac i i y sco es o speci ic ime poin s we e
highe a 20:00–04:00 h han a 10:00 h (ANOVA,
F
7,72
= 8.667, p< 0.001), bu du ing passi e win e ing
he mean alues did no di e a any ime poin s.
The e was a posi i e co ela ion be ween he ac i i y
sco es and T
b
o 9/9 indi iduals when simul aneous
ac i i y and T
b
alues we e included in he analysis
(
s
= 0.348–0.611, n = 162–780, p< 0.001), and in 7/9
animals when 24-h sums o ac i i y sco es and a e -
age 24-h T
b
alues we e co ela ed (
s
= 0.339–0.767,
n=26–129, p< 0.05). The a e age 24-h ac i i y sco e
had posi i e co a iance wi h he a e age 24-h T
a
(ANCOVA, F
1,6.645
= 11.019, p< 0.05) and nega i e co-
a iance wi h pho ope iod (F
1,510.584
= 9.288, p<0.01) and
dep h o snow (F
1,11.384
= 5.077, p< 0.05; Figu e 3), while
he e was no signi ican co a iance be ween he ac i i y
and dep h o so snow.
12/01 01/01 02/01 03/01 04/01 05/01 06/01
34
35
36
37
38
39
Body empe a u e
Da e (2009-10)
Figu e 1 Passi e pe iods o win e ing de e mined by change-poin analysis om body empe a u e da a. The body empe a u e (°C) o
an adul male accoon dog (M9) was measu ed e e y 120 min wi h in a-abdominal da a logge s in win e 2009–2010, ed line = ac i e pe iods,
blue line = passi e win e ing.
Mus onen e al. BMC Ecology 2012, 12:27 Page 6 o 17
h p://www.biomedcen al.com/1472-6785/12/27
Die a y analysis and a y acid signa u es
The s omachs had no diges ible ma e ial in 15 (16%) and
he in es ines in 5 cases (5%). The o al mass o he con-
en s a ied be ween 0–987 g. The mass was he highes
in No (287 ± 48 g), dec eased be ween No and Jan,
was he lowes in Feb (13 ± 4 g), and inc eased be ween
Feb and Ap il (K uskal–Wallis ANOVA, H= 34.303, d
=5,p< 0.001). The a e age olumes o all diges ible ood
i ems we e 71 ± 11 ml (s omachs) and 21 ± 3 ml (in es-
ines; see Addi ional ile 1). Undiges ible, man-made ma-
e ial (pieces o pape , plas ic, ciga e e bu s, e c.) was
ound in 27% o he s omachs and 23% o he in es ines.
Roundwo ms we e p esen in 29% o he indi iduals
wi h highe occu ence in No –Jan compa ed o Feb–
Ap il (χ
2
- es , χ
2
= 4.750, d = 1, p< 0.05).
Based on he FO1–2, olumes, and RS%, he impo -
ance o he main g oups o ood i ems dec eased as ol-
lows: mammals ≥plan s ≥bi ds ≥ ish ≥in e eb a es (see
Addi ional ile 2, Addi ional ile 3, Addi ional ile 4,
Addi ional ile 5, Addi ional ile 6, Addi ional ile 7,
Addi ional ile 8, Addi ional ile 9). Twel e mammals
could be classi ied o he species le el in he s omachs
and 10 in he in es ines. Roden s we e he mos com-
mon ood i ems among mammals (see Addi ional ile 2,
Rela i e equency
Ampli ude
0.0
0.1
0.2
0.3
0.4
0.5
0.6
Rela i e e
q
uenc
y
Ampli ude
0.0
0.1
0.2
0.3
0.4
0.5
0.6
24 h
12 h 8 h
24 h
12 h 8 h 6 h
a
b
Figu e 2 Rep esen a i e body empe a u e ampli ude spec a
du ing o e win e ing. The body empe a u e ampli ude spec a o
an adul male accoon dog (M1) du ing (a) ac i e and (b) passi e
pe iods o win e ing.
Mon h (2007-8)
Decembe Janua y Feb ua y Ma ch Ap il
Ac i i y sco es (AU), snow dep h (cm)
0
50
100
150
200
250
300
350
Da
y
len
g
h (h)
0
3
6
9
12
15
Ac i i y sco es
Snow dep h
Day leng h
Mon h (2007-8)
Decembe Janua y Feb ua y Ma ch Ap il
Ambien em
p
e a u e (ºC)
-20
-10
0
10
20
30
40
Body empe a u e (ºC)
34
35
36
37
38
39
Ambien empe a u e
Body empe a u e o M6
a
b
Figu e 3 A e age 24-h body empe a u es and ac i i y sco e
sums and hei ela ion o ex e nal ac o s. (a) The a e age 24-h
body empe a u es (°C) and (b) noc u nal ac i i y sco e sums
(a bi a y uni s; AU) o a ju enile male accoon dog (M6) we e
measu ed in Dec 2007–Ma ch 2008 oge he wi h he a e age 24-h
ambien empe a u es (°C), day leng h (h), and snow dep h (cm).
Mus onen e al. BMC Ecology 2012, 12:27 Page 7 o 17
h p://www.biomedcen al.com/1472-6785/12/27
Addi ional ile 3, and Addi ional ile 9). They consis ed
mainly o bank oles (Myodes gla eolus), ield oles
(Mic o us ag es is), and uniden i ied Mic o us oles, bu
he e we e also se e al obse a ions o ed squi els
(Sciu us ulga is). Roden s we e accompanied by insec-
i o es (almos exclusi ely So ex spp. sh ews), ha es
(Lepus spp.), ca ni o es (almos exclusi ely accoon
dogs), and ce ids (Ce idae) wi h ela i ely equal
occu ences. The e we e no di e ences in he FO1–2,
olumes, o RS% o hese ood i ems be ween he s o-
machs and in es ines.
Twen y edible plan s could be classi ied o he species
le el in he s omachs and 18 plan s in he in es ines (see
Addi ional ile 4, Addi ional ile 5, and Addi ional ile 9).
Ce eals we e he mos commonly u ilized plan -based
ood ollowed by be ies, ege ables, and ui s. A ena
sa i a was he mos impo an ce eal assessed by FO1–2
and olume, and he gas oin es inal ac s o some indi i-
duals con ained also small olumes o Panicum miliaceum.
The mos common be ies we e So bus aucupa ia,
Vaccinium i is-idaea,andV. oxycoccos bu hei olumes
we e gene ally small. The mos p e alen ege able species
we e Solanum ube osum and Daucus ca o a, whe eas
Py us communis,Malus domes ica,andMusa sp. we e
he mos common ui s. O he impo an plan s we e
Helian hus annuus by FO1–2 and olume and A achis
hypogaea by olume. The s omachs had highe olumes
o plan s han he in es ines (Mann–Whi ney U- es , U=
1044.000, n = 45, 61, p< 0.05). The s omachs had also
highe olumes o use ul plan s (Mann–Whi ney U- es ,
U= 947.500, n = 44, 60, p< 0.05) bu hei occu ence was
lowe (χ
2
- es , χ
2
=5.583,d =1,p<0.05).
The iden i ied bi ds we e o 6 species and 6 amilies,
bu e en he mos common a ian emains (Phasianidae
and Co idae) we e qui e a e and ound in he gas o-
in es inal ac s o only 5 animals (see Addi ional ile 6).
The s omachs had highe occu ence o bi ds han he
in es ines (χ
2
- es , χ
2
= 6.751, d = 1, p< 0.01). The con-
sumed ish we e iden i ied as belonging o 3 species and
4 amilies; he mos common amily was Pe cidae ol-
lowed by Cyp inidae and Esocidae (see Addi ional ile 7).
The in e eb a es we e almos exclusi ely insec s ha
could no be classi ied o he species le el, and hei
olumes we e e y small, highe in he s omachs han in
he in es ines (Mann–Whi ney U- es , U= 14.500, n = 8,
9, p< 0.05; see Addi ional ile 8).
The mos oluminous ood ypes in he s omachs we e
bi ds (12.3% o he cases), oa (9.9%), accoon dogs
(9.9%), ha es (9.9%), and bank oles (9.9%), and in he
in es ines, oa (13.6%), sh ews (13.6%), ha es (9.7%),
oles/lemmings (9.7%), and accoon dogs (7.8%). The s o-
machs o he emales had highe Vol1 (Mann–Whi ney
U- es , U= 795.500, n = 45, 48, p< 0.05) and highe occu -
ences o accoon dogs, medium-sized mammals, bi ds,
and ce eals (χ
2
- es , χ
2
=4.019–8.689, d = 1, p<0.01–
0.05) han hose o he males, which showed highe RS%
o bank oles and o al small mammals (Mann–Whi ney
U- es , U= 6.000, 137.000, n = 5–26, p< 0.05). The di e -
si y index was highe in he s omachs in Ap il compa ed
o Jan–Feb (K uskal–Wallis ANOVA, H= 13.653, d = 5,
p< 0.05). In ea ly win e , he accoon dogs consumed
mo e mammals and oa (K uskal–Wallis ANOVA, H=
7.138–14.737, d = 2, p<0.001–0.05) and less ish (s om-
ach: K uskal–Wallis ANOVA, H=7.977, d =2, p<0.05).
Vol1–2 we e he highes in No –Dec and dec eased in
Jan–Feb (K uskal–Wallis ANOVA, H= 15.202–21.445,
d = 5, p< 0.001–0.01).
The accoon dogs wi h ish in hei gas oin es inal
ac s a he ime o sampling had highe p opo ions o
pa icula n-3 polyunsa u a ed FA (PUFA; 22:4n-3,
22:5n-3, DHA 22:6n-3), o al n-6 PUFA (LA 18:2n-6,
22:4n-6, 22:5n-6), C20–24 sa u a ed FA (SFA), and
many C20–22 monounsa u a ed FA (MUFA) in he sc
a , while he pe cen ages o o al MUFA (mainly 18:1n-
9) we e lowe han in he animals wi hou ish ( - es ,
| | = 2.082–4.114; Mann–Whi ney U- es , U= 62.500–
119.000; n = 10, 41, p< 0.001–0.05). The p esence o
bi ds was associa ed wi h inc eased p opo ions o some
C14–17 SFA, a highe n-3/n-6 PUFA a io, and a
dec eased 20:1n-9 pe cen age ( - es , | | = 2.153–2.529,
n = 14, 15, p< 0.05). In PCA, he FAS o he animals,
whose gas oin es inal ac s con ained ish, we e sepa-
a ed om hose wi hou ish bu he e was no clea dis-
c imina ion acco ding o he o he die a y i ems. The
ela i e p opo ions o mos C12–17 SFA, pa icula
C14–17 MUFA, and C18–20 n-3 PUFA dec eased du -
ing win e while he pe cen ages o mos C18–22 SFA
and MUFA oge he wi h mos C20–22 n-6 PUFA and
C21–22 n-3 PUFA inc eased ( - es , | | = 2.071–8.625;
Mann–Whi ney U- es , U= 628.500–645.500; n = 39, 46,
p< 0.001–0.05). The p opo ions o o al MUFA and n-6
PUFA inc eased and hose o o al SFA and n-3 PUFA
dec eased. The p e e ence o FA mobiliza ion can be
seen in Addi ional ile 10.
Home ange sizes and habi a p e e ences
Fo each animal, 56–208 eloca ions we e ob ained du -
ing he s udy pe iods. The p opo ion o success ul GPS
posi ioning a emp s a ied om 16 o 62%. The inc e-
men al analysis sugges ed ha eliable home ange es i-
ma es we e achie ed wi h app oxima ely 50–140 ixes.
Eigh a ea-obse a ion cu es app oached he asymp o e
indica ing s able home anges. The a e age home ange
size was 5.3 ± 1.1 km
2
(K95%) and he size o co e a eas
1.3 ± 0.3 km
2
(K50%; Table 1). The alues we e highe
when a ju enile male (M5) wi h a e y la ge home ange
ha expanded ac oss a lake was included in he analyses.
Two ju eniles pe o med long dispe sions om hei
Mus onen e al. BMC Ecology 2012, 12:27 Page 8 o 17
h p://www.biomedcen al.com/1472-6785/12/27
ini ial home anges o new a eas: he sum o dis ances
be ween successi e eloca ions was 12 km o F2 (11 km
as he s aigh -line dis ance) and 60 km o M5 (33 km).
The yea o he s udy o he sex o he animals did no
a ec he sizes o home anges. In con as , he ju eniles
had la ge home anges han he adul s (K95% wi h M5:
16.0 ± 8.6 s. 3.5 ± 1.3 km
2
, Mann–Whi ney U- es , U=
5.000, n = 6, p< 0.05).
The p opo ional a ailabili y o habi a s was as ollows:
ields (ce eal ields, pas u es, allow a mland; 26% o he
main habi a s), coni e ous o es s on mine al soil (17%),
mixed o es s on mine al soil (13%), spa sely o es ed
a eas (canopy co e <30% and/o heigh <5 m; 13%), ice
(lakes, ponds, i e s, di ches; 11%), deciduous o es s on
mine al soil (7%), sho es (lakes, ponds, i e s, di ches;
4%), and ga dens (including ya ds; 4%). The mos com-
monly u ilized habi a s included spa sely co e ed a eas
(20.7 ± 6.5% o he eloca ions), ields (18.3 ± 3.0%), de-
ciduous o es s on mine al soil (13.8 ± 2.5%), ga dens
(11.0 ± 1.8%), mixed o es s on mine al soil (8.7 ± 1.5%),
and sho es (7.6 ± 2.4%; Figu e 4a). When he u iliza ion
o bio opes and passages was p opo ioned o hei a ail-
abili y in he a ea ha he animals could ha e used (1 =
equal a ailabili y and use, <1 = a oidance, >1 = p e e -
ence), he ollowing p e e ence was ob ained: ga dens
(3.4 ± 0.7), oads and oadsides (3.2 ± 0.7), sho es (2.5 ±
0.7), deciduous o es s on mine al soil (2.1 ± 0.5),
spa sely o es ed a eas (1.4 ± 0.4), ail oads and ail oad
beds (1.4 ± 1.0), ields (1.1 ± 0.3), mixed o es s on min-
e al soil (0.8 ± 0.2), coni e ous o es s on mine al soil
(0.4 ± 0.1), and ice (0.2 ± 0.1; Figu e 4b). The yea o he
s udy o he age o he animals did no a ec he esul s,
bu he emales equen ed coni e ous o es s on mine al
soil sligh ly less han he males did ( - es , | | = 2.338,
n=6,p< 0.05).
Gene al a iables
The a e age BM was he highes in Dec, eached he
nadi in Ma ch, and inc eased be ween Ma ch and
Ap il (ANOVA, F
5,78
= 17.577, p< 0.001). The BMI
dec eased signi ican ly be ween Jan and Feb, and he
mass o omen al a and he hickness o en al sc
a dec eased om Feb o Ma ch (ANOVA, F
5,58–87
=
7.953–15.535, p< 0.001). The au umnal BMI did no
co ela e signi ican ly wi h he numbe o passi e pe -
iods o win e ing, he o al sum o passi e days, o
he du a ions o he sho es and longes passi e pe i-
ods. The blood hemoglobin concen a ion was sligh ly
highe and he mean co puscula olume lowe in
sp ing compa ed o au umn (Mann–Whi ney U- es ,
U= 10.500, 9.000, n = 6, 10, p<0.05), bu he o he
pa ame e s in he CBC we e no in luenced by season
(da a no shown).
The a e age win e ime T
a
, calcula ed o he pe iod
wi h pe manen snow co e , was –7.2 ± 0.3°C (minimum
–29.0°C, maximum +7.0°C) in 2006–2007, –3.4 ± 0.2°C
(–15.0°C, +5.5°C) in 2007–2008, –6.4 ± 0.2°C (–23.0°C,
+2.5°C) in 2008–2009, and –11.5 ± 0.3°C (–32.0°C, +5.5°C)
in 2009–2010. The maximum snow dep hs on cul i a ed
land we e 45, 70, 41, and 82 cm, espec i ely.
Discussion
De e mina ion o win e sleep pe iods wi h change-poin
analysis
Following he ac i i y pa e ns o noc u nal and sec e i e
wild mammals is o en logis ically di icul . GPS acking
is eplacing con en ional adio acking and isual obse -
a ions o , e.g., snow acks as he p incipal me hod o
moni o ing win e ac i i y, bu all hese p ocedu es a e
economically challenging (expensi e, labou -in ensi e,
e c.). We de eloped he e a new low-cos , low-e o ap-
plica ion o CPA o de e mine he iming and du a ion
o al e na ing ac i e and passi e pe iods o o e win e ing
using he accoon dog as he model species. CPA has
been sca cely used in physiological esea ch e en hough
some s udies ha e u ilized his me hod [27,28]. The
p esen expe imen is he i s o show ha CPA can be
success ully applied o analyze T
b
eco dings o a pas-
si ely win e ing wild mammal in o de o un a el i s o -
aging pa e ns wi h po en ial applica ions o o he
no he n ca ni o es.
The posi i e co ela ion be ween he ac i i y sco es
and T
b
o he accoon dogs gi es an oppo uni y o d aw
conclusions on he iming o passi e pe iods by using
he T
b
da a alone. To alida e he me hod, he pe iods o
passi i y we e de e mined by bo h beha io al da a and
Table 1 Sizes o win e home anges o wild accoon
dogs (n = 12) in eas e n Finland
ID Reloca ions (n) Ke nel
95% (km
2
)
Ke nel
50% (km
2
)
Range
span (km)
M1 189 2.0 0.5 2.5
M2 66 10.0 1.6 8.5
M3 188 9.5 2.4 7.1
M4 104 1.0 0.2 3.2
M5 127 58.6 16.9 14.0
M6 208 2.4 0.6 3.3
F1 170 5.6 1.9 4.8
F2
a
59 10.5 2.3 6.3
F3 103 0.6 0.1 2.4
F4 197 4.5 1.5 5.0
F5 92 9.0 2.4 7.6
F6 56 3.3 1.1 3.6
Mean ± SE
b
130 ± 18 5.3 ± 1.14 1.3 ± 0.26 4.9 ± 0.65
M = male, F = emale,
a
he mean o wo home anges,
b
M5 wi h a e y la ge
home ange excluded.
Mus onen e al. BMC Ecology 2012, 12:27 Page 9 o 17
h p://www.biomedcen al.com/1472-6785/12/27
Depa men o Biology, Uni e si y o Oulu, P.O. Box 3000, FI-90014, Oulu,
Finland.
7
Finnish Game and Fishe ies Resea ch Ins i u e, P.O. Box 413,
FI-90014, Oulu, Finland.
8
Finnish Game and Fishe ies Resea ch Ins i u e,
I äinen Pi käka u 3, FI-20520, Tu ku, Finland.
9
Depa men o Biology,
Uni e si y o Tu ku, FI-20014, Tu ku, Finland.
10
Municipal Ve e ina y Clinic o
Joensuu, Takila ie 5, FI-80110, Joensuu, Finland.
Recei ed: 12 June 2012 Accep ed: 6 Decembe 2012
Published: 13 Decembe 2012
Re e ences
1. Rodge s AR: T acking animals wi h GPS: he i s 10 yea s. In T acking
Animals wi h GPS. An In e na ional Con e ence held a he Macaulay Land Use
Resea ch Ins i u e: 12–13 Ma ch 2001. Edi ed by Sibbald AM, Go don IJ:
Abe deen; 2001:1–10.
2. Kauhala K, Holmala K, Sch egel J: Seasonal ac i i y pa e ns and
mo emen s o he accoon dog, a ec o o diseases and pa asi es,
in sou he n Finland. Mamm Biol 2007, 72:342–353.
3. Mikkola M: Managemen Plan o he Raccoon Dog. Helsinki: Suomen
iis akeskus; 2011. In Finnish.
4. Melis C, He indal I, Kauhala K, Ande sen R, Høgda K-A: P edic ing animal
pe o mance h ough clima ic and plan phenology a iables: he case
o an omni o e hibe na ing species in Finland. Mamm Biol 2010,
75:151–159.
5. Mus onen A-M, Asikainen J, Kauhala K, Paakkonen T, Nieminen P: Seasonal
hy hms o body empe a u e in he ee- anging accoon dog
(Nyc e eu es p ocyonoides) wi h special emphasis on win e sleep.
Ch onobiol In 2007, 24:1095–1107.
6. Ki ao N, Fukui D, Hashimo o M, Osbo ne PG: O e win e ing s a egy o
wild ee- anging and enclosu e-housed Japanese accoon dogs
(Nyc e eu es p ocyonoides albus). In J Biome eo ol 2009, 53:159–165.
7. an Ma ken Lich enbel WD, Daanen HAM, Wou e s L, F onczek R, Raymann
RJEM, Se e ens NMW, Van Some en EJW: E alua ion o wi eless
de e mina ion o skin empe a u e using iBu ons. Physiol Beha 2006,
88:489–497.
8. Nieminen P, Saa ela S, Pyykönen T, Asikainen J, Mononen J, Mus onen A-M:
Endoc ine esponse o as ing in he o e win e ing cap i e accoon dog
(Nyc e eu es p ocyonoides). J Exp Zool 2004, 301A:919–929.
9. Kauhala K, Helle E: Age de e mina ion o he accoon dog in Finland.
Ac a The iol 1990, 35:321–329.
10. McNay RS, Mo gan JA, Bunnell FL: Cha ac e izing independence o
obse a ions in mo emen s o Columbian black- ailed dee . J Wildl
Manage 1994, 58:422–429.
11. Rooney SM, Wol e A, Hayden TJ: Au oco ela ed da a in eleme y s udies:
ime o independence and he p oblem o beha iou al e ec s. Mammal
Re 1998, 28:89–98.
12. de Solla SR, Bondu iansky R, B ooks RJ: Elimina ing au oco ela ion educes
biological ele ance o home ange es ima es. J Anim Ecol 1999,
68:221–234.
13. Wo on BJ: Ke nel me hods o es ima ing he u iliza ion dis ibu ion in
home- ange s udies. Ecology 1989, 70:164–168.
14. Kenwa d RE, Sou h AB, Walls SS: Ranges6 1.2: Fo he Analysis o T acking
and Loca ion Da a. Wa eham: Ana ack L d; 2003.
15. Kauhala K, Helle E, Taskinen K: Home ange o he accoon dog
(Nyc e eu es p ocyonoides) in sou he n Finland. J Zool 1993, 231:95–106.
16. Sii onen L, Sulka a S: Pohjolan nisäkkää (Mammals o No he n Eu ope).
Helsinki: O a a; 1994. In Finnish.
17. Cappe s RTJ, Bekke RM, Jans JEA: Digi ale Zadena las an Nede land.
G oningen: Ba khuis Publishing & G oningen Uni e si y Lib a y; 2006.
18. Ko ela M, F eyho J: Handbook o Eu opean F eshwa e Fishes. Co nol:
Publica ions Ko ela ; 2007.
19. Oa es DW, K ings LM, Di z KL: Field manual o he iden i ica ion o selec ed
No h Ame ican eshwa e ish by ille s and scales. Neb aska Technical Se ies
No. 19. Lincoln: Neb aska Game and Pa ks Commission; 1993. A ailable
online a [h p://icwdm.o g/inspec ion/Fish/FishManual.pd ]. Accessed on
Ma ch 25 2012.
20. Sido o ich VE, Solo ej IA, Sido o ich AA, Dyman AA: Seasonal and annual
a ia ion in he die o he accoon dog Nyc e eu es p ocyonoides in
no he n Bela us: he ole o habi a ype and amily g oup. Ac a The iol
2008, 53:27–38.
21. Day MG: Iden i ica ion o hai and ea he emains in he gu and aeces
o s oa s and weasels. J Zool 1966, 148:201–217.
22. Deed ick DW, Koch SL: Mic oscopy o hai Pa II: a p ac ical guide and
manual o animal hai s. Fo ensic Science Communica ions 2004, 6, a ailable
online a [h p://www. bi.go /abou -us/lab/ o ensic-science-
communica ions/ sc/july2004/ esea ch/2004_03_ esea ch02.h m]. Accessed
on Jan 19 2012.
23. Mus onen A-M, Asikainen J, Aho J, Nieminen P: Selec i e seasonal a y
acid accumula ion and mobiliza ion in he wild accoon dog (Nyc e eu es
p ocyonoides). Lipids 2007, 42:1155–1167.
24. Taylo WA: Change-poin analysis: A powe ul new ool o de ec ing changes.
[h p://www. a ia ion.com/cpa/ ech/changepoin .h ml]. Accessed on Feb 3
2012.
25. Taylo WA: A pa e n es o dis inguishing be ween au o eg essi e and mean-
shi da a. [h p://www. a ia ion.com/cpa/ ech/pa e n.h ml]. Accessed on
Feb 3 2012.
26. K alheim OM, Ka s ang TV: A gene al-pu pose p og am o mul i a ia e
da a analysis. Chemome In ell Lab 1987, 2:235–237.
27. Acke man JT, Takekawa JY, K use KL, O hmeye DL, Yee JL, Ely CR, Wa d
DH, Bollinge KS, Mulcahy DM: Using adio eleme y o moni o ca diac
esponse o ee-li ing ule g ea e whi e- on ed geese (Anse albi ons
elgasi) o human dis u bance. Wilson Bulle in 2004, 116:146–151.
28. Cassidy M, Mazzone P, Oli ie o A, Insola A, Tonali P, Di Lazza o V, B own P:
Mo emen - ela ed changes in synch oniza ion in he human basal
ganglia. B ain 2002, 125:1235–1246.
29. Tanaka H: Win e hibe na ion and body empe a u e luc ua ion in he
Japanese badge , Meles meles anakuma.Zool Sci 2006, 23:991–997.
30. Kowalczyk R, Jęd zejewska B, Zalewski A: Annual and ci cadian ac i i y
pa e ns o badge s (Meles meles) in Białowieża p ime al o es (eas e n
Poland) compa ed wi h o he Palaea c ic popula ions. J Biogeog aphy
2003, 30:463–472.
31. Hissa R, Siekkinen J, Hoh ola E, Saa ela S, Hakala A, Pudas J: Seasonal
pa e ns in he physiology o he Eu opean b own bea (U sus a c os
a c os) in Finland. Comp Biochem Physiol 1994, 109A:781–791.
32. Fowle PA, Racey PA: O e win e ing s a egies o he badge , Meles meles,
a 57 °N. J Zool 1988, 214:635–651.
33. Ha low HJ: To po and o he physiological adap a ions o he badge
(Taxidea axus) o cold en i onmen s. Physiol Zool 1981, 54:267–275.
34. Hwang YT, La i iè e S, Messie F: Ene ge ic consequences and ecological
signi icance o he e o he my and social he mo egula ion in s iped
skunks (Mephi is mephi is). Physiol Biochem Zool 2007, 80:138–145.
35. Aleksiuk M, S ewa AP: Food in ake, weigh changes and ac i i y o
con ined s iped skunks (Mephi is mephi is) in win e . Am Midland Na
1977, 98:331–342.
36. Be ange K, B øse h H: Body empe a u e changes in wild-li ing badge s
Meles meles h ough he win e . Wildli e Biol 1998, 4:97–101.
37. Kanda LL, Fulle TK, F iedland KD: Tempe a u e senso e alua ion o
opossum win e ac i i y. Wildl Soc Bull 2005, 33:1425–1431.
38. Kauhala K, Kaunis o M, Helle E: Die o he accoon dog, Nyc e eu es
p ocyonoides, in Finland. Z Säuge ie kunde 1993, 58:129–136.
39. Kobylińska J: The ed ox and accoon dog in we lands o he Bieb za
i e alley — ood composi ion and bu ow use. J Wildl Res 1996,
1:186–189.
40. Sido o ich VE, Polozo AG, Lauzhel GO, K asko DA: Die a y o e lap among
gene alis ca ni o es in ela ion o he impac o he in oduced accoon
dog Nyc e eu es p ocyonoides on na i e p eda o s in no he n Bela us.
Z Säuge ie kunde 2000, 65:271–285.
41. Bal ūnai ėL: Die composi ion o he ed ox (Vulpes ulpes L.), pine
ma en (Ma es ma es L.) and accoon dog (Nyc e eu es p ocyonoides
G ay) in clay plain landscape, Li huania. Ac a Zool Li uanica 2002,
12:362–368.
42. Bal ūnai ėL: Die and win e habi a use o he ed ox, pine ma en and
accoon dog in Dzūkija Na ional Pa k, Li huania. Ac a Zool Li uanica 2006,
16:46–53.
43. Pe ke ičius S, Nansen P, S ephenson L: The e ec o as ing on Asca is
suum and Oesophagos omum spp. In g owing pigs. In J Pa asi ol 1997,
27:431–437.
Mus onen e al. BMC Ecology 2012, 12:27 Page 16 o 17
h p://www.biomedcen al.com/1472-6785/12/27

44. Vi o P, Mikkola H: Food composi ion o he accoon dog Nyc e eu es
p ocyonoides G ay, 1834 in Finland. Z Säuge ie kunde 1981, 46:20–26.
45. Sasaki H, Kawaba a M: Food habi s o he accoon dog Nyc e eu es
p ocyonoides i e inus in a moun ainous a ea o Japan. J Mamm Soc
Japan 1994, 19:1–8.
46. Kauhala K, Laukkanen P, on Rége I: Summe ood composi ion and ood
niche o e lap o he accoon dog, ed ox and badge in Finland.
Ecog aphy 1998, 21:457–463.
47. Su o A, Kauhala K, Anso ge H: Die o he accoon dog Nyc e eu es
p ocyonoides –a canid wi h an oppo unis ic o aging s a egy.
Ac a The iol 2010, 55:165–176.
48. Fed iani JM, Fulle TK, Sau ajo RM: Does a ailabili y o an h opogenic
ood enhance densi ies o omni o ous mammals? An example wi h
coyo es in sou he n Cali o nia. Ecog aphy 2001, 24:325–331.
49. Kim C-H, Lee C-G, Yoon H-C, Nam H-M, Pa k C-K, Lee J-C, Kang M-I, Wee S-
H: Rabies, an eme ging disease in Ko ea. J Ve Med B 2006, 53:111–115.
50. Reig S, Jęd zejewski W: Win e and ea ly sp ing ood o some ca ni o es in
he Białowieża Na ional Pa k, eas e n Poland. Ac a The iol 1988, 33:57–65.
51. Jęd zejewski W, Jęd zejewska B, Szymu a A: Food niche o e laps in a
win e communi y o p eda o s in he Białowieża p ime al o es ,
Poland. Ac a The iol 1989, 34:487–496.
52. Sel a N, Jed zejewska B, Jed zejewski W, Waj ak A: Sca enging on
Eu opean bison ca casses in Bialowieza p ime al o es (eas e n Poland).
Écoscience 2003, 10:303–311.
53. Tanhuanpää E, Pulliainen E: Majo a y acid composi ion o some o gan
a s in he moose (Alces alces) in no heas e n Lapland. Ann Zool Fennici
1975, 12:148–155.
54. A o A, An oine JM, Pizzo e a o L, Reykdal O, an Poppel G: T ans a y
acids in dai y and mea p oduc s om 14 Eu opean coun ies: he
TRANSFAIR s udy. J Food Compos Anal 1998, 11:150–160.
55. Con esse P, Hegglin D, Gloo S, Bon adina F, Deplazes P: The die o u ban
oxes (Vulpes ulpes) and he a ailabili y o an h opogenic ood in he
ci y o Zu ich, Swi ze land. Mamm Biol 2004, 69:81–95.
56. Kauhala K: Habi a use o accoon dogs, Nyc e eu es p ocyonoides,in
sou he n Finland. Z Säuge ie kunde 1996, 61:269–275.
57. S e ens W: E ec s o a ia ion in essen ial a y acids in ish eeds on
nu i i e alue o eshwa e ish o humans. Aquacul u e 1997,
151:97–119.
58. Kousso oplis A-M, Lema chand C, Bec A, Des ile es C, Ambla d C, Fou nie
C, Be ny P, Bou die G: F om aqua ic o e es ial ood webs: dec ease o
he docosahexaenoic acid/linoleic acid a io. Lipids 2008, 43:461–466.
59. Zalewski K, Ma ysiak-Żu owska D, Iwaniuk M, Ni kiewicz B, S ołyhwo A:
Cha ac e iza ion o a y acid composi ion in Eu asian badge (Meles
meles). Polish J En i on S ud 2007, 16:645–650.
60. Kauhala K, Holmala K: Op imal adio- acking s a egy – he bes esul s
wi h he leas e o ? Ac a The iol 2008, 53:333–341.
61. Holmala K: The communi y o medium-sized ca ni o es: he in e ac ions
be ween species, habi a s and abies. PhD hesis: Uni e si y o Helsinki,
Depa men o Biological and En i onmen al Sciences; 2009.
62. Holmala K, Kauhala K: Habi a use o medium-sized ca ni o es in
sou heas Finland —key habi a s o abies sp ead? Ann Zool Fennici
2009, 46:233–246.
63. Kauhala K, Au ila M: Habi a p e e ences o he na i e badge and he
in asi e accoon dog in sou he n Finland. Ac a The iol 2010, 55:231–240.
64. Recio MR, Ma hieu R, Maloney R, Seddon PJ: Fi s esul s o e al ca s (Felis
ca us) moni o ed wi h GPS colla s in New Zealand. New Zeal J Ecol 2010,
34:288–296.
65. Med i IM, Mou ão G: Home ange o gian an ea e s (My mecophaga
idac yla) in he Pan anal we land, B azil. J Zool 2005, 266:365–375.
66. D ygala F, Zolle H, S ie N, Ro h M: Dispe sal o he accoon dog
Nyc e eu es p ocyonoides in o a newly in aded a ea in Cen al Eu ope.
Wildl Biol 2010, 16:150–161.
67. Åhlén P-A, Dahl F: Må dhunda u e på lång and ing. S ensk Jak 2011, 3
(issue):16. In Swedish.
68. Takeuchi T, Ma suki R, Nashimo o M: GPS cell phone acking in he
G ea e Tokyo A ea: a ield es on accoon dogs. U ban Ecosys 2012,
15:181–193.
69. D ygala F, S ie N, Zolle H, Boegelsack K, Mix HM, Ro h M: Habi a use o
he accon dog (Nyc e eu es p ocyonoides) in no h-eas e n Ge many.
Mamm Biol 2008, 73:371–378.
70. P ange S, Geh SD, Wigge s EP: In luences o an h opogenic esou ces on
accoon (P ocyon lo o ) mo emen s and spa ial dis ibu ion. J Mammal
2004, 85:483–490.
71. Bal ūnai ėL: Win e habi a use, niche b ead h and o e lap be ween he
ed ox, pine ma en and accoon dog in di e en landscapes o
Li huania. Folia Zool 2010, 59:278–284.
72. Seile A: Ecological E ec s o Roads, a Re iew. In oduc o y Resea ch Essay no
9. Uppsala: Swedish Uni e si y o Ag icul u al Sciences, Depa men o
Conse a ion Biology; 2001.
73. McLellan BN, Shackle on DM: G izzly bea s and esou ce-ex ac ion
indus ies: e ec s o oads on beha iou , habi a use and demog aphy.
J Appl Ecol 1988, 25:451–460.
74. Van Dyke FG, B ocke RH, Shaw HG, Acke man BB, Hemke TP, Lindzey FG:
Reac ions o moun ain lions o logging and human ac i i y. J Wildl
Manage 1986, 50:95–102.
75. Klein DR: Reac ion o eindee o obs uc ions and dis u bances. Science
1971, 173:393–398.
76. Inouye DW, Ba B, A mi age KB, Inouye BD: Clima e change is a ec ing
al i udinal mig an s and hibe na ing species. P oc Na l Acad Sci 2000,
97:1630–1633.
77. Ma ila V, G anholm H, Laanika i J, Y jölä T, Aal o A, Heikinheimo P,
Honka ukia J, Jä inen H, Liski J, Me i i a R, Paunio M: Finland’s Na ional
S a egy o Adap a ion o Clima e Change. Helsinki: Minis y o Ag icul u e
and Fo es y; 2005.
doi:10.1186/1472-6785-12-27
Ci e his a icle as: Mus onen e al.:Applica ion o change-poin analysis
o de e mine win e sleep pa e ns o he accoon dog (Nyc e eu es
p ocyonoides) om body empe a u e eco dings and a mul i- ace ed
die a y and beha io al s udy o win e ing. BMC Ecology 2012 12:27.
Submi you nex manusc ip o BioMed Cen al
and ake ull ad an age o :
• Con enien online submission
• Tho ough pee e iew
• No space cons ain s o colo figu e cha ges
• Immedia e publica ion on accep ance
• Inclusion in PubMed, CAS, Scopus and Google Schola
• Resea ch which is eely a ailable o edis ibu ion
Submi you manusc ip a
www.biomedcen al.com/submi
Mus onen e al. BMC Ecology 2012, 12:27 Page 17 o 17
h p://www.biomedcen al.com/1472-6785/12/27