1
Seasonal mo emen & ac i i y
o Akia-Manii soq ca ibou cows
in Wes G eenland
as de e mined by sa elli e
Technical Repo No. 99, 2017
G eenland Ins i u e o Na u al Resou ces
2
Ti le: Seasonal mo emen and ac i i y o Akia-Manii soq ca ibou
cows in Wes G eenland as de e mined by sa elli e
Au ho s: Ch is ine Cuyle 1, John Nagy2 and Ka l Zingle sen1
1G eenland Ins i u e o Na u al Resou ces, P.O. Box 570,
3900–Nuuk, G eenland
2CW405 Biological Science Cen e , Uni e si y o Albe a,
Albe a, Canada
Se ies: Technical Repo No. 99, 2017
Da e o publica ion: 19 July, 2017
Publishe /Funding: Pinngo i ale i ik, G eenland Ins i u e o Na u al Resou ces
Co e pho o: Polled Akia-Manii soq ca ibou cow, Augus : pho o C. Cuyle
ISBN: 87-91214-76-9
ISSN: 1397-3657
EAN: 9788791214769
Ci ed as: Cuyle , C., Nagy, J. & Zingle sen, K. 2017. Seasonal
mo emen and ac i i y o Akia-Manii soq ca ibou cows in
Wes G eenland as de e mined by sa elli e.
Pinngo i ale i ik – G eenland Ins i u e o Na u al
Resou ces. Technical Repo No. 99. 94 pp.
Con ac add ess: The epo is only a ailable in elec onic o ma . You can
download a PDF- ile o he epo a his homepage
h p://www.na u .gl/publika ione / ekniske appo e
I is possible o ob ain a p in o he epo he e:
G eenland Ins i u e o Na u al Resou ces
P.O. Box 570
DK-3900 Nuuk
G eenland
Phone: +299 36 12 00
Fax: +299 36 12
E-mail: in o@na u .gl
www.na u .gl
3
Seasonal mo emen s & ac i i y
o
Akia-Manii soq ca ibou cows
in Wes G eenland
as de e mined by sa elli e
By
Ch is ine Cuyle 1, John Nagy2 & Ka l Zingle sen1
1 G eenland Ins i u e o Na u al Resou ces, P.O. Box 570, 3900–Nuuk, G eenland
2 CW405 Biological Science Cen e , Uni e si y o Albe a, Albe a, Canada
Technical Repo No. 99, 2017
G eenland Ins i u e o Na u al Resou ces
4
[Emp y page]
5
Table o Con en s
Summa y (English) ................................................................... 8
Eqikkaaneq (G eenlandic) ..................................................... 10
Resume (Danish) ..................................................................... 12
In oduc ion ............................................................................. 15
Me hods .................................................................................... 19
Resul s ...................................................................................... 27
Discussion ................................................................................ 49
Acknowledgemen s ................................................................. 64
Li e a u e ci ed ........................................................................ 65
Lis o Figu es
1.
Map o en i e Akia-Manii soq egion. Ele a ions abo e 200 me e s a e
ligh yellow while hose below a e g een.
Page 16
2.
Akia-Manii soq s udy a ea, 11,969 km2, p e iously known cal ing a eas
Page 17
3.
The 26 mo ali ies among sa elli e-colla ed ca ibou cows, e c.
Page 28
4.
O e iew o mo emen pa hs o sa elli e-colla ed Akia-Manii soq ca ibou
cows 2008-2010.
Page 30
5.
Mean daily mo emen by Julian day o Akia-Manii soq ca ibou cows
Page 31
6.
Dis ibu ion o habi a ypes p esen wi hin 150x150 m mo ing window
a ound each ocal 30 m g id cell in he s udy a ea, e c.
Page 32
7.
Ca ego ized p obabili y o occu ence o Akia-Manii soq ca ibou cows in
Wes G eenland du ing 10 ac i i y pe iods, e c.
Page 41
8.
Ele a ion o si es used du ing seasonal ac i i y pe iods by Akia-Manii soq
ca ibou cows in Wes G eenland, 2008-2010.
Page 42
9.
Wes G eenland Akia-Manii soq sa elli e-colla ed ca ibou, p obabili y o
cow occu ence du ing peak cal ing, 21 May o 8 June: e c.
Page 43
10.
Dis ibu ion o he 52 cal ing si es 2008-2010 in ela ion o subs a e, o
he sa elli e-colla ed Akia-Manii soq ca ibou cows Wes G eenland, e c.
Page 44
11.
A: Daily mo emen pa e n o pa u ien cow: e c.
B: Daily mo emen pa e n o non-pa u ien cow: e c.
Page 46
12.
Annual a ia ion in he numbe o ca ibou cows in a ious ep oduc i e
s a es, 2008-2010.
Page 48
13.
Tempo al dis ibu ion o 52 pa u i ion da es 2008-2010 o sa elli e-
colla ed Akia-Manii soq ca ibou cows, Wes G eenland, e c.
Page 49
14.
Sa elli e colla deploymen loca ions, 40 ca ibou cows, 1-7 May 2008, e c.
Page 75
15.
Ca ibou mo emen ou es obse ed, 1-7 May 2008, e c.
Page 77
6
Lis o Tables
1.
Labels & de ini ions o esou ce selec ion (RS) habi a and ele a ion
co a ia es conside ed in RS models, e c.
Page 23
2.
Mo ali y among sa elli e-colla ed Akia-Manii soq ca ibou cows, e c.
Page 27
3.
Mean and o al numbe o loca ions ob ained o acked emales and
ac i i y pe iods, espec i ely, 2008-2010.
Page 27
4.
Seasonal ac i i y pe iods as calcula ed om daily a el a es, e c.
Page 30
5.
A ailabili y o habi a ypes wi hin 0.02 km2 a ea a ound ocal g id cells
Page 31
6.
Bes - i ac i i y pe iod esou ce selec ion unc ion models, e c.
Page 33
7.
Pa ame e es ima es & s anda d e o s in bes - i cal ing and peak o
cal ing (mean cal ing da e ±1 SD) ac i i y pe iod RSF models, e c.
Page 34
8.
Pa ame e es ima es & s anda d e o s in bes - i pos -cal ing, ea ly/mid-
summe , and mid/la e summe ac i i y pe iod RSF models, e c.
Page 35
9.
Pa ame e es ima es & s anda d e o s in bes - i all/p e-b eeding,
b eeding, and pos -b eeding/la e all ac i i y pe iod RSF models, e c.
Page 36
10.
Pa ame e es ima es & s anda d e o s in bes - i ea ly/mid-win e , la e
win e , and p e-cal ing ac i i y pe iod RSF models, e c.
Page 37
11.
Habi a selec ion by seasonal ac i i y pe iod, e c.
Page 38
12.
Habi a selec ion du ing he cal ing pe iod ela i e o hose selec ed
du ing all o he ac i i y pe iods, e c.
Page 39
13.
Pe cen a ailabili y, pe cen use, & ‘ a io o % use:% a ailabili y’ o a eas
by p obabili y o occu ence ca ego y and ac i i y pe iod, e c.
Page 40
14.
Compa ison o cal ing si e condi ions chosen by adul cows (n=30)
ela i e o p esence o absence o a yea ling-a -heel, e c.
Page 45
15.
Compa ison o cal ing si e condi ions chosen by adul cows (n =51; sub
adul emo ed) 2008-2010, Akia-Manii soq Wes G eenland
Page 45
16.
Sa elli e colla deploymen posi ions on 40 ca ibou cows, e c.
Page 76
17.
Cal ing ca ibou esou ce selec ion models
Page 78
18.
Peak-cal ing ca ibou esou ce selec ion models
Page 79
19.
Pos -cal ing ca ibou esou ce selec ion models
Page 80
20.
Ea ly summe ca ibou esou ce selec ion models
Page 81
21.
Mid /la e summe ca ibou esou ce selec ion models
Page 82
22.
Fall /p e-b eeding ca ibou esou ce selec ion models
Page 83
23.
B eeding ( u ) ca ibou esou ce selec ion models
Page 84
24.
Pos -b eeding /la e all ca ibou esou ce selec ion models
Page 85
25.
Ea ly/mid-win e ac i i y pe iod ca ibou esou ce selec ion models
Page 86
26.
La e win e ac i i y pe iod ca ibou esou ce selec ion models
Page 87
27.
P e-cal ing ac i i y pe iod ca ibou esou ce selec ion models
Page 88
28.
Cal ing e sus pos -cal ing, ea ly summe & mid /la e summe , e c
Page 89
29.
Cal ing e sus all/p e-b eeding, b eeding & pos -b eeding/la e all, e c
Page 90
30.
Cal ing e sus ea ly/mid-win e , la e-win e , and p e-cal ing, e c.
Page 91
31.
Rep oduc i e s a us o colla ed cows, e c.
Page 92
7
Lis o Appendices
1.
Cap u e o Akia-Manii soq ca ibou cows, 1-7 May 2008.
Page 75
2.
SBIC (Schwa z-Bayesian in o ma ion c i e ion) di e ences o 10 bes - i
esou ce selec ion models o each ac i i y pe iod.
Page 78
3.
Habi a selec ion by G eenland ca ibou (Rangi e a andus g oenlandicus)
du ing he CALVING pe iod ela i e o o he ac i i y pe iods, e c
Page 89
4.
Rep oduc i e s a us o colla ed cows ali e a cal ing
Page 92
5.
Equipmen ecommenda ions o sa elli e colla ing
Page 93
*Raw da a a ailable a G eenland Ins i u e o Na u al Resou ces:
F:/40-59/PaFu/42/Landpa edy / 35 ALCOA Rensdy spo ing
8
Summa y (English)
In May 2008 GPS sa elli e colla s we e deployed on 40 cows om he Akia-
Manii soq (AM) ca ibou (Rangi e a andus g oenlandicus) popula ion. In he pe iod
2008-2010, he sa elli e-colla ed cows p o ided loca ions, which we analysed o
mo emen s, spa ial dis ibu ion, cal ing si es, loca ion a ibu es, seasonal ac i i y
pe iods, and habi a esou ce selec ion. The AM ca ibou a e subs an ially less
documen ed han he ds in No h Ame ica. The esul s in his epo a e pa icula ly
aluable o managemen in G eenland because hey es ablish a baseline o AM
habi a use in he cu en absence o signi ican de elopmen and in as uc u e in
he Cen al egion. Howe e , he sho s udy pe iod and small sample size,
exace ba ed by high mo ali y, weaken esul s. Al hough ob aining colla da a in
G eenland is di icul and cos ly, a longe ime se ies wi h a s able la ge numbe o
colla ed ca ibou is necessa y be o e sweeping conclusions can be suppo ed.
AM cows beha e simila ly o he moun ain eco ype o ca ibou a he han he
ba en-g ound. Cal ing was no con ined o speci ic exclusi e cal ing g ounds close
o he Ice Cap. Bi hing cows spaced-away in a con inuum ac oss he en i e Cen al
egion, om seacoas o Ice Cap. The e is li le ele ance o p o ec ing speci ic
exclusi e cal ing g ounds in he AM egion. Ins ead, conse a ion measu es would
p o i om applying a b oad-scale habi a managemen app oach o he widesp ead
AM cal ing habi a . Ele a ion was a good p edic o o p obabili y o cow occu ence
a cal ing. Whe he in a xe ic zone o no , sou h acing slopes a ound 600 m ele a ion
wi h snow we e a ou ed. Al hough iming and wid h o he eme gen ege a ion
pe iod and possibly ain a oidance may be he d i ing ac o s behind choice o high
ele a ion o cal ing by pa u ien AM cows, hese emain o be in es iga ed. AM
cal ing appea s o begin ea lie han p e iously assumed, and sugges s ha he
pe iod o p o ec ion measu es migh be shi ed o wa d. Habi a possessing he
p e e ed a ibu es o bi hing comp ises 42-45% o he Cen al egion. Thus
al hough cal ing ange is essen ial o ca ibou p oduc ion, a p esen i is no likely
limi ing he AM popula ion. Howe e , gi en he high ideli y by AM cows o
p e ious bi hing si es, a wa ming A c ic o an h opogenic in luences could ha e
nega i e impac s i cows a e displaced o habi a s less a ou able o cal su i al.
Mo ali y among sa elli e-colla ed cows was high and much appea ed due o
ha es . I common o he en i e popula ion, his would ha e played a majo ole in
he decline o AM abundance om 2001 o 2010.
Pa e ns o annual mo emen con i med a sou hwes -no heas axis. Dis ance
mo ed could be sho , wi h indi iduals ypically a he wes e n end o hei axis in
win e and eas e n end in summe . Each cow u ilized jus a ac ion o he a ailable
a ea. This s ongly indica es popula ion sub-di ision wi hin he Cen al egion and
sugges s ha he en i e AM popula ion will no be in luenced when ei he s ochas ic
wea he e en s o managemen ac ions a ec only a po ion o he egion.
Pa e ns o seasonal mo emen e ealed 10 seasonal ac i i y pe iods, which
we e associa ed wi h speci ic habi a a ibu es. Ele a ion was he p ima y habi a
9
a ibu e ha a ied signi ican ly ac oss he seasonal ac i i y pe iods. B eeding
occu ed a low ele a ions. I hun ing seasons coincide wi h he u , hen human
dis u bance may nega i ely in luence b eeding and subsequen ly cal p oduc ion.
Ca ibou ulne abili y o ha es would also likely inc ease because o hei ela i e
accessibili y o hun e s.
Daily mo emen pa e ns a ied h oughou he yea . Cows mo ed leas in
ea ly Ma ch making i op imal o ae ial su ey. July had maximum mo emen ,
sugges ing insec ha assmen as he cause and suppo ing he necessi y o insec
elie habi a . Du ing he cal ing pe iod, a cha ac e is ic daily mo emen pa e n was
assumed o indica e a bi hing e en , i.e., ise om no mal immedia ely p eceding a
sha p d op o nea ze o and he ea e a g adual ise. In u u e, bi hing could be
alida ed by isually loca ing sa elli e-colla ed cows by ai plane o helicop e , o
equipping he sa elli e-colla s wi h ideo capabili y.
Habi a a ibu es a e no e enly dis ibu ed. Rela i e o he en i e Cen al
egion, and in con as o cal ing habi a , he a ailable a ea is small o la e summe ,
all and win e habi a s. These could be limi ing o he AM popula ion. Speci ically,
he la ges ac o win e habi a , Akia, albei s ill small in size, is ulne able o sou h
wes e ly s o m sys ems ha can ende win e o age una ailable o ene ge ically
cos ly o access. Thus, special a en ion and p o ec ion h ough ine-scale habi a
managemen may be app op ia e. AM ca ibou abundance would likely bene i i o
hose habi a s ha a e sca ce, a) ca ibou access was p ese ed, b) an h opogenic
dis u bance was mi iga ed, and c) ha densi ies o AM ca ibou we e kep below
ca ying capaci y o hese limi ed anges.
Managemen is bes ailo -made o he popula ion, he seasonal ac i i y unde
conside a ion and he amoun o habi a a ailable o ha ac i i y. Conse a ion
e o s should add ess no jus one se e al seasonal anges i al o ep oduc ion,
insec elie and su i al. P o ec ing pa u ien cows and hei bi hing habi a is no
a one-sho cu e o ensu ing eco e y o sus ainabili y o ca ibou popula ions.
Ca ibou oam. Globally, ca ibou ange shi s a e common, e en o ‘sac osanc ’
cal ing a eas. Thus, managemen mus conside conse ing cu en ly unused a eas
o po en ial u u e use by ca ibou. Meanwhile, human in luences on he landscape
a e ecognized ac o s ha can exace ba e ca ibou declines. In No h Ame ica, he
cu en h eshold p oposed o p e en ing ca ibou decline, is ha 65% o he o al
ange emains unexposed o human dis u bance. P oac i e managemen and
conse a ion di ec ed owa ds p ese ing la ge undis u bed in ac landscapes,
ele an o se e al seasonal ac i i ies and hei mo emen co ido s, would os e
ca ibou conse a ion now and o u u e gene a ions.
16
a o able condi ions o cal su i al (Whi e 1983, Fancy & Whi en 1991).
Cal ing s a egies aim a maximizing su i al o new o sp ing, and may
include shi s in la i ude o ele a ion o educe p eda ion isk, ep oduc i e
synch ony o achie e p eda o sa ia ion, o p io i ize access o eme gen ood
esou ces and g een-up (Be ge ud & Page 1987, F yxell e al. 1988, Klein 1990,
Albon & Lang a n 1992, Pos e al. 2003, Loe e al. 2005). Cal ing phenology is
gene ally synch onous wi hin popula ions and closely ela ed o pho ope iod
as well as he onse and du a ion o plan g ow h (Pos 2003). I is also
a ec ed by leng h and ha shness o he p e ious win e (T e aa e al. 2013).
Since ca ibou a e ‘capi al’ b eede s, i.e., pa u ien cows depend on body
ese es a he han o age, peak pa u i ion can p ecede g een-up by se e al
weeks (Du an e al. 2005, Moen e al. 2006, Ba boza & Pa ke 2008, Taillon e
al. 2013). Because s a egies a y somewha e en among Rangi e ,
unde s anding he speci ic c i e ia ha cha ac e ize habi a i al o cal ing in
Wes G eenland is impo an o conse a ion measu es o ha e an in luence
on abundance.
Figu e 1. Map o en i e Akia-Manii soq egion, an a ea o ca. 15,400 km2. Ele a ions abo e 200 me e s
a e ligh yellow while hose below a e g een.
50°W
Manii soq
66°N
52°W
54°W
64°N
A ammik
Napasoq
NUUK
Kapisilli
G eenland
Ice Cap
Akia-Manii soq
Cen al egion
hun ing a ea 3
50 100
kilome es
0
G eenland
17
Al hough cen al Alaska and Yukon ca ibou ypically exhibi synch onous
and long 300-500 km no h-sou h mig a ions, wi h annual dis ances a elled
anging o 5000 km (C aighead & C aighead 1987, Fancy e al. 1989), Wes
G eenland ca ibou esemble he moun ain/ o es dwelling seden a y ca ibou
eco ype, because agg ega ions a e absen and mo emen s sho (50-60 km)
and non-synch onous (Cuyle & Linnell 2004). P e iously, hey we e assumed
o ha e wes -eas seasonal mig a ions be ween he seacoas (win e ) and
G eenland Ice Cap (summe ) when abundance was high, and sho ened
mig a ions o none a all i low (Vibe 1967, G ønnow e al. 1983, Thing 1984).
Ce ainly, G eenland ca ibou summe ange can be inland nea he Ice Cap;
howe e , win e ange can be in e media e be ween Ice Cap and seacoas
(Cuyle & Linnell 2004). Meanwhile, egionally ele an s udies a e lacking.
Figu e. 2. Akia-Manii soq s udy a ea, 11,969 km2, p e iously known cal ing a eas, ● Cuyle and
Linnell (2004), ● Cuyle unpublished, 2004 b own lines (Aas up and Nymand 2004), 2015 s ippled
b own (NunaGis 2015). Ele a ions below 200 m a e g een, abo e 200 m yellow.
Al hough AM ca ibou a e p esen h oughou he Cen al egion, cal ing
loca ions a e no adequa ely known. Th oughou No h Ame ica, ba en-
g ound ca ibou popula ions usually ha e one well iden i ied adi ional
cal ing g ound, a sub-a ea o he o al ange o a speci ic popula ion (Russell
e al. 2002). A cal ing g ound is a speci ic exclusi e a ea whe e 80-90% o
pa u ien cows e u n annually o cal e (Gunn & Mille 1986), bu may
18
change in spa ial loca ion o e ime (Gunn e al. 2008, Taillon e al. 2012).
Iden i ying cal ing g ounds o AM ca ibou has been elusi e; al hough
S andgaa d e al. (1983) sugges ed AM cal ing occu ed in he no he n
inland po ion o he ange. In he Kange lussuaq-Sisimiu ca ibou popula ion
o he no h, la ge cal ing cow agg ega ions in close p oximi y o he
G eenland Ice Cap we e documen ed by Thing (1984). Subsequen s udies o
G eenland cal ing g ounds ocused on inland a eas and obse ed cows
cal ing (Aas up 1986; Aas up & Nymand 2004; J. Nymand, G eenland
Ins i u e o Na u al Resou ces, unpublished da a). Meanwhile, hun e
knowledge o AM indica ed a small ugged highland a ea nea he AM
seacoas (200-500 m ele a ion; 65° 23’ N; 51° 40’ W). Fu he , eigh sa elli e-
colla ed AM cows, 1997-1999, we e highly dispe sed in he cal ing season
(Cuyle & Linnell 2004), no es ic ed o a eas close o he Ice Cap, we e
ypically a ele a ions abo e 300 m (Fig. 2) and exhibi ed 88% ideli y among
sequen ial bi hing si es (Cuyle & Linnell 2004). Si e ideli y (philopa y) is
he endency o an animal o s ay in, o habi ually e u n o, a speci ic si e o
a ea. These a ibu es a e simila o he moun ain ca ibou eco ype (Be ge ud
e al. 1984, Skogland 1989). Ne e heless, he e was jus one small inland
cal ing g ound o icially ecognized o p o ec ion (Aas up & Nymand 2004).
Al hough his cal ing g ound was expanded by 2015 (h p://www.
nunagis.gl), i emained a speci ic a ea associa ed wi h he pe iod 20 May o
20 June (Go e nmen o G eenland Bu eau o Mine als and Pe oleum 2000)
and human ac i i ies ha e been egula ed acco dingly.
Seasonal habi a s ha e been hi he o poo ly unde s ood and knowledge
insu icien o p o ide a i m basis o managemen ecommenda ions ha
can mee u u e challenges e.g., s ochas ic wea he e en s, in as uc u e
de elopmen s, pa hogens, o any combina ion o hese and o he ac o s.
P esen S udy
In ea ly May 2008, we deployed GPS sa elli e colla s on 40 ca ibou cows o he
AM popula ion and ollowed hese un il July 2010. We delinea ed mean daily
mo emen o e a 1-yea cycle and ex en o mig a o y pa h. We examined he
seasonal ac i i y pe iods o Akia-Manii soq ca ibou cows and desc ibe he
habi a a ibu es associa ed wi h hose ac i i y pe iods. We in es iga ed and
mapped he loca ion and a ailabili y o he habi a a ibu es and p obabili y
o ca ibou cow occu ence o a gi en ac i i y pe iod. We delinea e mo emen
pa e ns associa ed wi h cal ing si es, iming o cal ing pe iod, cal ing
loca ions, and cow ideli y o cal ing si es. We de e mined whe he a eas
19
used by ca ibou du ing cal ing ha e cha ac e is ics ha a e uniquely di e en
om hose used du ing es o he yea . En i onmen al condi ions we e
conside ed. Since adul emale su i al is a key de e minan o ungula e
popula ion dynamics, mo ali y among he sa elli e-colla ed cows is
add essed. Since secu ing su icien habi a could acili a e conse a ion o
AM ca ibou, we delinea e which seasonal anges a e bo h essen ial and
limi ed in a ailabili y.
Me hods
S udy a ea
The Cen al egion (hun ing a ea 3) o Wes G eenland is cen ed a 65° 10’ N;
51° 25’ W no h o Nuuk, and is ca. 15,400 km2. Ou s udy a ea, a sub-se o
he o al egion (Figs. 1, 2), was 11,969 km2 and excluded he hea ily glacia ed
no hwes , as well as lakes, glacie s and sand. The su ounding ocean, jo ds
and glacie s es ic dispe sal o ca ibou om he egion, c ea ing a la gely
"closed" popula ion. The egion is p ima ily ugged uplands and moun ains,
ele a ions ≥ 300 m co e 60%, 200-300 m co e 8% and unde 200 m 32%. The
egion is unde eloped and lacks in as uc u e. Du ing he s udy pe iod,
2008-2010, he backcoun y was inaccessible o he majo i y o people. Mos
we e limi ed o boa ing along he coas and jo d sho elines, and hiking in o
he e ain seldom was beyond abou 6 km. O he han ca ibou, mammals
p esen a e ew and include only he a c ic ha e (Lepus a c icus) and a c ic ox
(Alopex lagopus) as well as in oduced e al eindee (1952) (R. . a andus) and
he ecen ly a i ed muskoxen (O ibos moscha us; na u al immig a ion since
ca. 1998 om he Kange lussuaq popula ion no h o Sukke oppen Ice Cap).
La ge p eda o s a e absen ; howe e , he e is an annual ca ibou ha es . The
egion is sandwiched be ween a domina ing high p essu e o e he
G eenland Ice Cap o he eas and equen low-p essu e oceanic s o m
sys ems (wind speeds 22–46 m/sec) om he sou hwes (Tams o 2004, DMI
2014, Gambe g e al. 2016). P ecipi a ion dec eases s ongly wi h dis ance om
he seacoas , esul ing in a xe ic con inen al clima e nea he Ice Cap
(Tams o 2004). The nea es me eo ological s a ion is immedia ely sou h in
he seacoas ci y o Nuuk, which has an annual mean empe a u e o -1.4°C,
mean July 6.5°C and ecei es an annual p ecipi a ion o 752 mm (Tams o e
al. 2005, DMI 2014). Vege a ion is chie ly low-a c ic species. Plan
communi ies a y wi h ele a ion, aspec , and p oximi y o in luences o
G eenland's we coas al ma i ime and d y con inen al ice cap clima es (Lund
20
e al. 2004, Simonsen 2011). The Akia coas al lowlands consis p ima ily o
lichen- ich dwa sh ub hea h, which mo ing inland changes o dwa sh ub
hea h wi h inc easing g assland. In highe ele a ions windswep idges,
ab asion pla eaus and ba e g ound domina e (Tams o 2004). The dominan
plan species in ou ege a ion classes ollows Bay & Simonsen (2009).
De ailed desc ip ions o ege a ion a e in Lund e al. (2004) and Simonsen
(2011). Vege a ion and snow maps and digi al e ain models a e in Tø up
(2009).
Ca ibou cap u e & eleme y
Cow ca ibou we e cap u ed 1-7 May 2008 wi h a ne -gun i ed om a
helicop e . Sa elli e colla s we e deployed on 40 cows. The e we e 20 Telonics
(Mesa, A izona, USA and Se ice A gos, Lando e , Ma yland, USA) and 20
I idium Sys em Ne wo k (Vec onic Ae ospace GmbH, Be lin Ge many)
global posi ioning sys em (GPS) sa elli e colla s. In e -loca ion in e als o
Telonics and I idium colla s we e 1 hou (24 pe day) and 2 hou s (12 pe
day), espec i ely. All colla s we e p og ammed o elease au oma ically a e
108 -118 weeks. Loca ions om May 2008 o July 2010 we e eco ded as
longi ude and la i ude coo dina es and p ojec ed o he UTM Zone 22N
P ojec ed Coo dina e Sys em, WGS 1984 Geog aphic Coo dina e Sys em,
No he n Hemisphe e. Fo GIS analyses, we used A cMap 10.1
(En i onmen al Sys ems Resea ch Ins i u e, Inc., Redlands, Cali o nia, USA).
Ca ibou cap u e and colla ing loca ions we e sp ead h oughou he Cen al
egion sou h o Sukke oppen Ice Cap (Appendix 1). We iden i ied cows by
p esence o ul a pa ch. Cap u e was close o cal ing and in AM cows
abdominal swelling om win e umen diges a/con en s is insigni ican and
la e win e a ese es among p egnan cows a e gene ally ze o (Cuyle
unpublished da a om 1996, 1997, 2008). Viewed om abo e, cows we e
assumed p egnan i he sides o he pos e io hal o hei body p o uded
qui e ma kedly ela i e o o he cows. A cap u e, 38 o he 40 cows we e
asce ained p egnan (included a sub adul ). Too h e up ion and wea
de e mined age class (sub adul , adul ). Cows colla ed in May 2008 included,
39 adul s (age ≥ 36 mon hs) and one sub adul (35 mon hs). The o al numbe
o GPS loca ions ob ained in he 2008-2010 s udy pe iod was 262,137.
Daily mo emen s and mig a o y pa h leng h
We calcula ed he di ec line dis ance (km) be ween all sequen ial loca ions
pe day o each ca ibou cow. Daily a el a es we e s anda dized o he
numbe o possible loca ions pe day [sum o dis ances be ween loca ions
21
ob ained/(no. loca ions ob ained/no. loca ions possible pe day)] and
no malized using a log10 ans o ma ion. Calcula ed dis ances p o ide a
minimum es ima e o o al indi idual mo emen due o 1-2 hou gaps in
eleme y.
We mapped pa hs o illus a e mo emen a ia ion. All pa hs we e c ea ed
using Haw h’s ools (Beye 2007). We mapped he annual pa hs aken by each
ca ibou each yea . We summed daily pa h dis ances (km) o ob ain annual
mean o al pa h dis ance, maximum and minimum. Longi udes / la i udes o
he la e wo p o ided he leng h o a cow’s home ange. We gene a ed
annual (calcula ed om he da e o cap u e) minimum con ex polygons
(MCPs) and pa hs (s aigh -line dis ances be ween sequen ial loca ions) o
each ca ibou using Haw h’s Tools (Beye 2007). We measu ed MCP a eas and
pa h leng hs, s anda dized hese o 365 days (a eas o leng h di ided by
numbe o days acked x 365), and no malized hem using a log10
ans o ma ion. MCP a eas and pa h leng hs a e in luenced by sample size
(Bo ge e al. 2006).
Seasonal ac i i y pe iods
We subdi ided he calcula ed daily a el a e da a in o 73 5-day pe iods and
used analysis o a iance (ANOVA) and Tukey’s hones ly signi ican
di e ence (HSD) pai -wise compa isons (SPSS 11.5, Chicago, Illinois, USA) o
iden i y all sequen ial 5-day pe iods when mo emen a es we e no
signi ican ly di e en . These ga e he s a and end da es o each seasonal
ac i i y pe iod. We used he i s and las known and es ima ed pa u i ion
and concep ion da es o de ine he cal ing and b eeding pe iods espec i ely.
Habi a selec ion modelling
He bi o e mo emen a es (Nagy 2011) and habi a equi emen s o
main enance, g ow h, ep oduc ion (Fe guson & Elkie, 2004; Gus ine e al.,
2006; Ho n & Rubens ein, 1984) a y seasonally (Maie & Whi e, 1998).
Animal habi a a ini ies a e commonly quan i ied a landscape scales
(Johnson e al. 2004) using da a o a dependen a iable: eleme y animal
loca ions (use) and geog aphic in o ma ion sys em (GIS) gene a ed andom
loca ions (a ailable), i.e. 10 andom loca ions pe animal loca ion, wi h each
andom loca ion sampled om wi hin a ci cle ha was cen e ed on he
p eceding eleme y loca ion, and ha ing a adius equal o he dis ance
be ween he p eceding and nex successi e eleme y loca ion. The
independen p edic o a iables a e habi a and opog aphic ea u es hough
22
o in luence habi a selec ed by he ca ibou (Table 1). The da a o dependen
and independen a iables a e used o cons uc esou ce selec ion unc ions
(RSF) o he ollowing o m:
w(x) = exp (β1x1 + β2x2 + ... + βkxk) Equa ion 1
(Johnson e al. 2004). The ela i e p obabili ies o occu ence o animals on
landscapes a e commonly calcula ed om RSFs and mapped using linea
s e ch ans o ma ions o he o m:
ŵ = [(w(x) - wmin)/( wmax - wmin)] Equa ion 2
whe e w(x) is he p oduc o equa ion 1 and wmin and wmax ep esen he
smalles and la ges RSF alues, espec i ely (Johnson e al. 2004). Howe e ,
ela i e p obabili ies o occu ence assessmen s using his me hod a e
sensi i e o he alue o wmax. Ra e la ge alues o wmax may esul in unde -
es ima ion o he ela i e p obabili ies o occu ence and a ec he eliabili y
o k- old c oss- alida ion echniques ypically used o assess he p edic i e
pe o mance o RSF models (Boyce e al. 2002). The consequences o unde -
es ima ing habi a alues in an a ea may be signi ican o conse a ion and
managemen .
We assumed ha he a ibu es (e.g., ele a ion, ege a ion co e ypes, wind-
exposu e) o si es selec ed by G eenland ca ibou a ied seasonally, o add ess
hei changing physiological equi emen s and beha io s. The e o e, we i s
iden i ied hei ac i i y pe iods based on signi ican changes in daily
mo emen a es (Nagy 2011). We used a mo ing-window GIS app oach o
assess he p esence/absence o 11 ege a ion co e ypes and a e age
ele a ion wi hin a 0.02 km2 a ea a ound each ca ibou (use) and andom
(a ailable) loca ion. We used logis ic eg essions (equa ion 1) o i RSF
models o each ac i i y pe iod (Johnson e al. 2004, La ham e al. 2011) and
selec ed he bes - i ing models using Schwa z in o ma ion c i e ia (SBIC)
(Cook 2007, Ha din & Hilbe 2012, Schwa z 1978, S ong e al. 1999, Wasse e
al. 2011). To calcula e ela i e p obabili ies o ca ibou occu ence (w) (equa ion
3) we modi ied equa ion 2. Ra he han wmax we used he wmedian RSF alues
o ca ibou use loca ions wi hin he s udy a ea o each ac i i y pe iod using
he ollowing modi ied linea s e ch ans o ma ion:
ŵ = [(w(x) - wmin)/(wmedian RSF ca ibou loca ions - wmin)]. Equa ion 3
23
We used wmedian RSF ca ibou loca ions unde he assump ion ha a leas 50 pe cen
o he ca ibou loca ions ob ained du ing any ac i i y pe iod should be in
p e e ed habi a s. This me hod allows o eplicable mapping o RSF models
and compa isons o ela i e p obabili y o occu ence among seasonal
models. We pa i ioned ŵ in o h ee equal in e als o ela i e p obabili y o
occu ence ca ego ies: A) ≤0.333 (low), B) ˃ 0.333 and ≤ 0.666 (mode a e), and
C) ˃ 0.666 (high). We calcula ed he pe cen o loca ions ha ell wi hin each
p obabili y o occu ence ca ego y o quan i y he known equency o use o
a eas we mapped as high, mode a e, and low p obabili y o occu ence. We
ob ained ela i e p obabili y o occu ence o a speci ic ac i i y pe iod by
pooling da a o all yea s o ha ac i i y pe iod.
Table 1. Labels & de ini ions1 o esou ce selec ion (RS) habi a and ele a ion co a ia es conside ed in
seasonal RS models o Wes G eenland Akia-Manii soq sa elli e-colla ed ca ibou cows.
Co a ia e
De ini ion
Ele a ion
Con inuous a iable; maximum ele a ion in me e s abo e sea le el (m asl) wi hin
ocal a ea*.
Hea h
Disc e e a iable; p esen (1) o absen (0) wi hin ocal a ea. Vege a ion
domina ed by willow (Be ula nana), c owbe y (Empe um nig um
he maph odi um), bluebe y (Vaccinium uliginosum mic ophyllum), and Lab ado
ea (Ledum g oenlandicum, Ledum palus e decumbens).
Open hea h
Disc e e a iable; p esen (1) o absen (0) wi hin ocal a ea. Vege a ion
domina ed by bluebe y (Vaccinium uliginosum mic ophyllum), bi ch (Be ula
nana), hea he (Cassiope e agona, Phyllodoce coe ulea), and Lab ado ea
(Ledum palus e decumbens).
Copse
Disc e e a iable; p esen (1) o absen (0) wi hin ocal a ea. Vege a ion
domina ed by willow (Salix glauca).
Fen
Disc e e a iable; p esen (1) o absen (0) wi hin ocal a ea. Vege a ion
domina ed by i eweed (E ipho um angus i olium hypa c icum) and sedges
(Ca ex a i lo a).
G ass
Disc e e a iable; p esen (1) o absen (0) wi hin ocal a ea. Vege a ion
domina ed by g asses (Calamag os is lapponica/g oenlandica), sedges (Ca ex
b unnescens, Ca ex bigelowii), bunch g asses (Deschampsia lexuosa), and wood
ushes (Luzula spica a).
Snow-bed
Disc e e a iable; p esen (1) o absen (0) wi hin ocal a ea. Vege a ion
domina ed by willows (Salix he bacea) and sedges (Ca ex bigelowii).
Wind-exposed
Disc e e a iable; p esen (1) o absen (0) wi hin ocal a ea. Vege a ion
domina ed by mosses, lichens, dwa sh ubs (Diapensia lapponica lapponica),
Laba do ea (Ledum palus e decumbens), campion (Silene acaulis), swee g ass
(Hie ochloë alpine), sedges(Ca ex bigelowii), and hododend ons (Rhododend on
lapponica).
Soil / Rock
Disc e e a iable; p esen (1) o absen (0) wi hin ocal a ea. Exposed soil and
ock.
Sedimen
Disc e e a iable; p esen (1) o absen (0) wi hin ocal a ea. Ri e , es ua ine, o
coas al lood plains.
Wa e
Disc e e a iable p esen (1) o absen (0) wi hin ocal a ea. F esh wa e lakes,
i e s, and s eams.
Snow / Ice
Disc e e a iable p esen (1) o absen (0) wi hin ocal a ea. Pe sis en mul i-
annual snow and ice.
1 Bay & Simonsen 2009; Simonsen 2011
*Focal a ea is he 5x5 30 m cell mo ing window a ound each 30 m cell.
24
Model cons uc ion: animal loca ions & en i onmen al co a ia es
We de i ed he 12 independen a iables (also e med co a ia es) om
Tø up’s (2009) 30 m esolu ion sa elli e image based digi al habi a and
ele a ion models (Table 1), which included hea h, open-hea h, copse, en,
g ass, snow-bed, wind-exposed, soil o ock, sedimen , wa e , snow o ice
(Bay & Simonsen 2009; Simonsen 2011) and ele a ion. We used A cMap o
c ea e a sepa a e 30 m cell-size g id o each independen a iable. Habi a
g id cells we e assigned alues o 1 o 0 i habi a s we e p esen o absen ,
espec i ely. We used a mo ing window, 5x5 30 m cell, in GIS, o de e mine
he p esence (1) o absence (0) o habi a s wi hin an a ea o ≈ 0.02 km2
su ounding each 30 m g id cell ( ocal cell) o each habi a g id. Focal cells
we e eclassi ied as ha ing no da a i clouds o shadows occu ed in >20% o
he cells con ained in he mo ing window. We assigned he maximum
ele a ion wi hin mo ing windows o he ele a ion g id ocal cells. We used
GIS o ex ac he alue o each ocal habi a and ele a ion g id cell o
in e sec ca ibou and andom loca ions (gene a ed a a a e o 1 pe km2 in he
s udy a ea).
Resou ces selec ion analysis
Da a o each emale we e sub-sampled andomly o one loca ion pe day (o
he mos accu a e loca ions). 19,572 loca ions (one loca ion pe day pe emale)
we e used o calcula e esou ce selec ion unc ions. We assessed pa e ns o
habi a selec ion a he popula ion le el (Johnson e al. 2004; La ham e al.
2011), i.e., a he scale o he landscape wi hin he s udy a ea, by compa ing
habi a s and ele a ions a ca ibou GPS (used) and andom (a ailable)
loca ions wi h used- e sus-a ailable design logis ic eg essions o each
seasonal ac i i y pe iod (Johnson e al. 2006, Manly e al. 2002). We used GLM
logi STATA 9 (STATCORP, College S a ion, Texas, USA) o assess he
ela ionship be ween use and a ailabili y o all possible combina ions o he
12 explana o y a iables and selec ed he bes - i ing models (Cook 2007,
Ha din & Hilbe 2012, Schwa z 1978, S ong e al. 1999, Wasse e al. 2011) o
each seasonal ac i i y pe iod using Schwa z’s in o ma ion c i e ia (SBIC)
(STATA 9; Schwa z 1978). We mapped he p obabili y o occu ence o
ca ibou using ou modi ied linea s e ch ans o ma ion (equa ion 3).
Assessmen o habi a selec ion among ac i i y pe iods
We compa ed ela i e selec ion o habi a ypes by ca ibou du ing cal ing and
all o he ac i i y pe iods using logis ic eg essions o es ima e coe icien s o
la en selec ion di e ence (LSD) unc ions (Cze we ynski 2007, La ham e al.
25
2011, Muelle e al. 2004). We used ANOVA and Tukey’s hones ly signi ican
di e en (HSD) pai wise compa isons o de e mine i ele a ions a ca ibou
loca ions a ied signi ican ly among ac i i y pe iods. We log10 ans o med
ele a ion da a p io o conduc ing ANOVA and back- ans o med he
esul ing s a is ics o in e p e a ion.
Cal ing
Pa u i ion (cal ing si e): mo emen pa e ns
Ce ids commonly exhibi ma ked (i.e., > 50%) declines in daily mo emen s
immedia ely ollowing pa u i ion (Long e al. 2009). An ab up lack o
mo emen is e iden when ca ibou bi h hei cal ; cows s op comple ely
(Len 1966). Following bi h o he neona e, di ec ional mo emen is absen
o se e al hou s and he ea e ; i i occu s, i is slow wi h many pauses (Len
1966). Mo e ecen ly, Fe guson & Elkie (2004) obse ed ha a seden a y
pe iod o low mo emen a e main ained o abou 3 days a i med cal ing in
woodland ca ibou (Rangi e a andus ca ibou). Using his knowledge and daily
mo emen a es in he ±10 days a ound he da e a cow s opped comple ely
Nagy (2011) asce ained da e o bi hing o 336 cows, which was alida ed by
isual su ey. C i e ia included daily mo emen a es ising sha ply o
se e al days, hen alling p ecipi ously o nea ze o, wi h a g adual inc ease in
he days ollowing.
We p edic ed pa u i ion da es using he abo e. We calcula ed daily a el
a e by each ca ibou cow du ing he 15 Ap il - 15 July maximum ange o
po en ial cal ing da es, and assumed ha he da e o he ab up all/cessa ion
in daily mo emen signaled a bi hing e en , while he associa ed GPS
loca ion p o ided he cal ing si e poin da a.
Cal ing pe iod & cow ideli y o cal ing si e
In addi ion o mean cal ing da e, we de ined pe iods o ‘mos ’ and ‘peak’
cal ing o augmen assessmen o possible empo al a ia ions. The pe iod o
‘mos ’ cal ing was es ima ed o be wi hin ±1.96 s anda d de ia ion (SD; 95%
CI) o he mean pa u i ion da e, and ‘peak’ cal ing o be wi hin ±1 SD (68%
CI) (Nagy 2011). We es ima ed concep ion da es by back-da ing 229 days om
pa u i ion da es (Be ge ud 1975, McEwan & Whi ehead 1972, Rowell &
Shipka 2009). We used he i s and las es ima ed pa u i ion and concep ion
da es o de ine he s a and end da es o he cal ing and b eeding pe iods,
espec i ely. Al hough he numbe o colla ed cows declined s eeply o e he
s udy pe iod, we s ill a emp ed o examine whe he he pe iod o cal ing
32
Figu e 6. Dis ibu ion o habi a ypes p esen wi hin 150x150 m mo ing window a ound each ocal
30 m g id cell in he s udy a ea. Mul i-yea pe manen snow/ice a iable doesn’ include Ice Cap.
Use / A ailable a ios
The a ios indica e when ca ibou use o a habi a was disp opo iona ely less
han, equal o, o g ea e han a ailabili y wi hin he s udy a ea (<1.0, 1.0, and
>1.0). The highes UA a io was 3.33 du ing he la e all/ea ly win e season
(Table 13). Thus, cows we e disp opo iona ely selec ing o his ange in ligh
o i s low a ailabili y. The second highes UA was 3.03 and occu ed du ing
b eeding. The hi d was all/p e-b eeding a 2.75 ollowed closely by mid-
win e a 2.71. The ea e came la e win e and mid/la e summe , bo h wi h
UA’s >2.0. Gi en ha ea ly/mid-summe ange was he mos a ailable i was
no su p ising ha i also had he lowes UA, 1.49.
33
Table 6. Bes - i ac i i y pe iod esou ce selec ion unc ion models o Akia-Manii soq ca ibou cows in he Cen al egion o Wes G eenland,
2008-2010.
Ac i i y pe iod
Co a ia es in bes - i models
BIC
Cal ing
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-150298.7
Peak-cal ing
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-137189.4
Pos -cal ing
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-136853.8
Ea ly/mid-summe
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-140048.1
Mid/la e summe
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-140439.1
Fall/p e-b eeding
hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-137204.7
B eeding
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-143218.3
La e all/ea ly win e
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-154203.3
Mid-win e
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-144092.0
La e win e
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-139452.9
P e-cal ing
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-137443.7
34
Table 7. Pa ame e es ima es and s anda d e o s in he bes - i cal ing and peak o cal ing (mean cal ing da e ±1 SD) ac i i y pe iod esou ce selec ion
unc ion models o Akia-Manii soq ca ibou cows in he Cen al egion o Wes G eenland, 2008-2010.
Habi a
co a ia e
Cal ing Pe iod
(mean cal ing da e ±1.96 SD)
Peak cal ing
(mean cal ing da e ±1 SD)
Pa ame e
es ima e
SE
P- alue
Pa ame e
es ima e
SE
P- alue
Ele a ion
0.001
0.000
0.000
0.002
0.000
0.000
Hea h
0.487
0.057
0.000
0.294
0.084
0.000
Open-hea h
0.661
0.061
0.000
0.429
0.086
0.000
Copse
0.870
0.061
0.000
0.889
0.090
0.000
Fen
-0.103
0.058
0.074
0.051
0.083
0.540
G ass
0.446
0.047
0.000
0.401
0.068
0.000
Snow-bed
-0.210
0.104
0.044
-0.010
0.152
0.950
Wind exposed
0.431
0.058
0.000
0.350
0.084
0.000
Soil/ ock
-0.203
0.049
0.000
-0.223
0.072
0.000
Sedimen
-0.238
0.070
0.001
-0.383
0.106
0.000
Wa e
-0.605
0.074
0.000
-0.907
0.116
0.000
Snow/ice
-1.275
0.329
0.000
-2.295
0.715
0.000
In e cep
-3.050
0.091
0.000
-3.734
0.132
0.000
35
Table 8. Pa ame e es ima es and s anda d e o s in he bes - i pos -cal ing, ea ly/mid-summe , and mid/la e summe ac i i y pe iod esou ce selec ion unc ion
models o Akia-Manii soq ca ibou cows in he Cen al egion o Wes G eenland, 2008-2010.
Habi a
co a ia e
Ac i i y pe iod
Pos -cal ing
Ea ly/mid summe
Mid/la e summe
Pa ame e
es ima e
SE
P- alue
Pa ame e
es ima e
SE
P- alue
Pa ame e
es ima e
SE
P- alue
Ele a ion
0.002
0.000
0.00
0.001
0.000
0.00
0.001
0.000
0.00
Hea h
0.907
0.092
0.00
0.558
0.078
0.00
1.385
0.088
0.00
Open-hea h
-0.002
0.083
0.98
-0.249
0.066
0.00
-0.073
0.069
0.29
Copse
0.884
0.086
0.00
0.974
0.075
0.00
0.794
0.068
0.00
Fen
0.017
0.086
0.85
0.111
0.070
0.11
0.037
0.072
0.60
G ass
0.290
0.071
0.00
0.276
0.063
0.00
0.254
0.063
0.00
Snow-bed
0.235
0.142
0.10
0.380
0.104
0.00
0.289
0.103
0.01
Wind exposed
0.391
0.083
0.00
0.195
0.071
0.01
-0.321
0.070
0.00
Soil/ ock
-0.324
0.076
0.00
-0.127
0.065
0.05
0.096
0.065
0.14
Sedimen
-0.102
0.099
0.31
0.115
0.073
0.12
0.006
0.078
0.94
Wa e
-0.936
0.116
0.00
-0.749
0.084
0.00
-0.498
0.085
0.00
Snow/ice
-2.219
0.716
0.00
-1.260
0.329
0.00
-15.682
494.905
0.98
In e cep
-3.802
0.132
0.00
-2.903
0.104
0.00
-3.384
0.111
0.00
36
Table 9. Pa ame e es ima es and s anda d e o s in he bes - i all/p e-b eeding, b eeding, and pos -b eeding/la e all ac i i y pe iod esou ce selec ion unc ion models
o Akia-Manii soq ca ibou cows in he Cen al egion o Wes G eenland, 2008-2010.
Habi a
co a ia e
Ac i i y pe iod
Fall/p e-b eeding
B eeding
Pos -b eeding/la e all
Pa ame e
es ima e
SE
P- alue
Pa ame e
es ima e
SE
P- alue
Pa ame e
es ima e
SE
P- alue
Ele a ion
-
-
-
-0.002
0.000
0.00
-0.004
0.000
0.00
Hea h
1.189
0.093
0.00
1.258
0.096
0.00
1.257
0.082
0.00
Open-hea h
-
-
-
0.023
0.069
0.74
0.385
0.062
0.00
Copse
0.602
0.074
0.00
0.449
0.062
0.00
0.348
0.052
0.00
Fen
-0.155
0.089
0.08
-0.048
0.072
0.50
0.063
0.058
0.28
G ass
0.679
0.068
0.00
0.710
0.060
0.00
0.505
0.055
0.00
Snow-bed
-0.034
0.151
0.82
-0.288
0.128
0.03
-0.310
0.101
0.00
Wind exposed
-0.065
0.077
0.40
-0.391
0.070
0.00
-0.121
0.059
0.04
Soil/ ock
-0.172
0.076
0.02
-0.080
0.067
0.23
-0.452
0.060
0.00
Sedimen
-0.276
0.105
0.01
0.059
0.083
0.48
-0.040
0.069
0.56
Wa e
-0.555
0.112
0.00
-0.961
0.097
0.00
-0.656
0.076
0.00
Snow/ice
-14.893
510.544
0.98
-14.467
453.309
0.98
-
-
-
In e cep
-3.308
0.101
0.00
-2.253
0.111
0.00
-1.627
0.094
0.00
37
Table 10. Pa ame e es ima es and s anda d e o s in he bes - i ea ly/mid win e , la e win e , and p e-cal ing ac i i y pe iod esou ce selec ion models o Akia-Manii soq
ca ibou cows in he Cen al egion o Wes G eenland, 2008-2010.
Habi a
co a ia e
Ac i i y pe iod
Ea ly/mid win e
La e win e
P e-cal ing
Pa ame e
es ima e
SE
P- alue
Pa ame e
es ima e
SE
P- alue
Pa ame e
es ima e
SE
P- alue
Ele a ion
-0.003
0.000
0.00
-0.002
0.000
0.00
-0.001
0.000
0.00
Hea h
0.901
0.079
0.00
0.950
0.091
0.00
0.274
0.082
0.00
Open-hea h
0.653
0.076
0.00
0.304
0.079
0.00
0.529
0.087
0.00
Copse
0.397
0.061
0.00
0.505
0.069
0.00
0.435
0.079
0.00
Fen
0.132
0.063
0.04
-0.079
0.074
0.29
0.074
0.076
0.33
G ass
0.219
0.059
0.00
0.228
0.067
0.00
0.332
0.069
0.00
Snow-bed
-0.152
0.108
0.16
-0.408
0.133
0.00
-0.295
0.134
0.03
Wind exposed
0.383
0.063
0.00
0.348
0.073
0.00
0.446
0.077
0.00
Soil/ ock
-0.305
0.061
0.00
-0.190
0.070
0.01
-0.442
0.071
0.00
Sedimen
0.044
0.075
0.56
0.399
0.081
0.00
0.273
0.085
0.00
Wa e
-0.690
0.085
0.00
-0.661
0.095
0.00
-0.489
0.094
0.00
Snow/ice
-
-
-
-
-
-
-
-
-
In e cep
-2.352
0.103
0.00
-2.631
0.113
0.00
-2.734
0.115
0.00
38
Table 11. O e iew o habi a selec ion by seasonal ac i i y pe iod o Akia-Manii soq ca ibou cows in he Cen al egion o Wes G eenland, 2008-2010.
Seasonal
Ac i i y pe iod
Resou ce selec ion unc ion co a ia es*
ele a ion
hea h
open-hea h
copse
en
g ass
snow-bed
wind-exposed
soil/ ock
sedimen
wa e
snow/ice
cal ing
+
+
+
+
=
+
-
+
-
-
-
-
pos -cal ing
+
+
=
+
=
+
=
+
-
=
-
-
ea ly/mid summe
+
+
-
+
=
+
+
+
=
=
-
-
mid/la e summe
+
+
=
+
=
+
+
-
=
=
-
=
all/p e-b eeding
excluded
+
excluded
+
=
+
=
=
-
-
-
=
b eeding
-
+
=
+
=
+
-
-
=
=
-
=
pos -b eeding/la e all
-
+
+
+
=
+
-
-
-
=
-
excluded
ea ly/mid win e
-
+
+
+
+
+
=
+
-
=
-
excluded
la e win e
-
+
+
+
=
+
-
+
-
+
-
excluded
p e-cal ing
-
+
+
+
=
+
-
+
-
+
-
excluded
*+, =, and - deno e co a ia e coe icien s ha we e signi ican g ea e , no signi ican ly di e en om, and signi ican ly less han ze o.
39
Table 12. Habi a selec ion du ing he cal ing pe iod ela i e o hose selec ed du ing all o he ac i i y pe iods based on la en selec ion di e ence (LSD) unc ion
model compa isons o Akia-Manii soq ca ibou cows in he Cen al egion o Wes G eenland, 2008-2010.
Seasonal
Ac i i y Pe iod
Rela i e selec ion*
Hea h
Open-hea h
Copse
Fen
G ass
Snow-bed
Wind-exposed
Soil/ ock
Sedimen
Wa e
Snow/ice
Pos -cal ing
26
>100
=
=
=
31
=
=
=
>100
=
Ea ly summe
=
>100
=
18
>100
=
>100
=
31
=
=
Mid/la e summe
48
>100
=
=
=
39
>100
30
28
=
Fall/p e-b eeding
51
>100
=
=
20
=
>100
=
=
=
B eeding
68
>100
=
=
17
=
>100
=
43
>100
Pos -b eeding/la e all
74
=
>100
29
=
=
>100
>100
46
>100
=
Ea ly/mid win e
63
30
=
35
>100
=
>100
>100
47
>100
=
La e win e
62
=
=
17
>100
=
>100
=
59
>100
62
P e-cal ing
=
=
>100
24
>100
=
>100
>100
49
=
=
*Rela i e selec ion was calcula ed o a iables wi h coe icien s signi ican ly di e en om 0 as exp (b) when b>0 and as [1-exp(b) when b<0 (La ham, La ham & Boyce,
2011). Rela i e selec ion alues <100 indica e ha use o he habi a du ing cal ing was signi ican ly less by x% han ha o he season being compa ed, alues >100
indica e use o habi a du ing cal ing was signi ican ly g ea e han ha o he season being compa ed, and = indica es habi a use du ing cal ing and season being
compa ed we e no signi ican ly di e en (Appendix 3).
40
Table 13. Pe cen a ailabili y, pe cen use, and a io o pe cen use: pe cen a ailabili y o a eas by p obabili y o occu ence ca ego y and ac i i y pe iod o Akia-
Manii soq ca ibou cows in he Cen al egion o Wes G eenland, 2008-2010.
Seasonal
Ac i i y pe iod
P obabili y o occu ence
Pe cen o s udy a ea
by occu ence ca ego y
(A ailable)
Pe cen o ca ibou loca ions
by occu ence ca ego y
(Use)
U:A a io
o occu ence ca ego ies*
Low
Mode a e
High
Low
Mode a e
High
Low
Mode a e
High
Cal ing
29
26
45
5
16
79
0.17
0.62
1.77
Pos -cal ing
30
31
39
6
22
72
0.18
0.69
1.89
Ea ly/mid summe
17
31
52
3
20
77
0.19
0.65
1.49
Mid/la e summe
51
15
34
17
11
72
0.33
0.73
2.09
Fall/p e-b eeding
57
22
21
16
26
58
0.28
1.18
2.75
B eeding
63
13
24
13
13
74
0.21
1.01
3.03
La e all/ea ly win e
67
12
21
12
19
69
0.18
1.56
3.33
Mid-win e
57
17
26
13
16
71
0.23
0.94
2.71
La e win e
53
16
31
13
12
75
0.24
0.75
2.44
P e-cal ing
33
23
44
8
19
73
0.25
0.82
1.67
*U:A a ios <1.0, 1.0, and ˃1.0 indica e use was disp opo iona ely less han a ailable, p opo ional o a ailabili y, and disp opo iona ely g ea e han a ailabili y.
41
Figu e 7. Ca ego ized p obabili y o occu ence o Akia-Manii soq ca ibou cows in Wes G eenland du ing 10 ac i i y pe iods: ollow bold black a ows o annual
cycle om cal ing o p e-cal ing. Maps ep esen he p oduc o landscape-scale esou ce selec ion unc ions.
48
Mean Julian cal ing da es o adul cows 2008, 2009, and 2010 we e
espec i ely, 152.80 ±6.48 days ( ange 134-163; n = 30), 151.29 ±10.45 ( ange
132-165; n = 14) and 142.86 ±5.24 days ( ange133-149; n = 7). Mean cal ing
da es did no di e o 2008 and 2009, 2009 and 2010 (ANOVA F1 = 0.348, P =
0.558; F1 = 3.98, P = 0.061 espec i ely), howe e , a di e ence was obse ed
be ween 2008 and 2010 (ANOVA F1 = 14.193, P = 0.006). Employing he non-
pa ame ic Mann-Whi ney U es on cal ing da es esul ed in no di e ences
be ween 2008 and 2009, 2009 and 2010 (P > 0.05), bu di e ence be ween 2008
and 2010 (P ≤ 0.001). 2010 cal ing was app ox. 10 days ea lie han in 2008.
Mean Julian cal ing da es o only hose cows cal ing in he wo speci ic
yea s es ed also es ed non-signi ican o 2008 and 2009, 2009 and 2010 (P >
0.05). Compa ison o cows bi hing in bo h 2008 and 2010 was con ounded by
he small sample size o h ee and indi idual esul s o 15, 6 and -1 days,
whe e 2010 cal ing was a mean 7 days ea lie han in 2008 (P > 0.05).
Cal ing si e ideli y
Al hough sample size was small and he s udy pe iod sho , we no ed ha
cal ing si e ideli y was high. The e we e 16 sequen ial cal ing loca ions o
14 cows ( wel e cows had wo loca ions; wo cows had h ee). Median
dis ance be ween sequen ial cal ing loca ions was 7.1 km. Mean dis ance was
12.5 km ± 11.7 SD, minimum was 0.5 km, and maximum 41.5 km. The
minimum coincided wi h he cow ha cal ed in 2008 and again in 2010, a e
a hia us o one yea .
Figu e 12. Annual a ia ion in he numbe o ca ibou cows in a ious ep oduc i e s a es, 2008-2010.
0
5
10
15
20
25
30
35
2008 2009 2010
Numbe o cows
Yea
Cal
Unknown
No cal
49
Figu e 13. Tempo al dis ibu ion o 52 pa u i ion da es 2008-2010, Wes G eenland Akia-Manii soq
sa elli e-colla ed ca ibou cows (no e 28 June sub adul ou lie ).
Discussion
The AM ca ibou a e subs an ially less documen ed han ca ibou popula ions
in No h Ame ica. The e o e, his epo could bene i ca ibou managemen in
G eenland. Iden i ying he a ibu es, loca ions and ex en o ca ibou seasonal
habi a s, and he iming o use o hese, is indispensable o decisions in ended
o p edic and o se nega i e impac s o ca ibou habi a , e.g., by clima e and
an h opogenic ac o s. The esul s p esen ed in his epo a e pa icula ly
aluable because hey es ablish a baseline o AM habi a use in he cu en
absence o signi ican de elopmen and in as uc u e in he Cen al egion.
We do no ad ise ex apola ing he cu en knowledge om AM o o he
popula ions in G eenland, owing o di e ences in popula ion size, la i ude,
clima e, opog aphy and lack o seasonal habi a use da a o he o he
popula ions. Fu he , he sho s udy pe iod and small sample size, which
cons an ly diminished o e ha pe iod, weaken his s udy’s indings.
Akia-Manii soq cow mo emen s
Daily
Simila o Cuyle & Linnell (2004) his s udy obse ed minimum daily
mo emen /ac i i y du ing a ca. 30-day pe iod concu en wi h Ma ch,
speci ically a he beginning o Ma ch. Highes cow mo emen a es occu ed
in July, and may be associa ed wi h insec ha assmen . The au umn u
0
1
2
3
4
5
6
12/May 19/May 26/May 2/Jun 9/Jun 16/Jun 23/Jun
Numbe o pa u i ion da es
Da e
50
(b eeding) c ea ed a second b ie e peak in ac i i y in la e Sep embe . This
knowledge indica es ha ae ial su eys o ca ibou abundance in ea ly Ma ch
will minimize he p oblem o ca ibou mo emen a ec ing he esul ing
popula ion es ima es.
The ac i i y end obse ed is no new. A summe ac i i y inc ease has been
obse ed o ca ibou / eindee by se e al au ho s (Ban ield 1954, Segal 1962,
Thomson 1971, Gaa e e al. 1975, Whi e e al. 1975, and Roby 1977). The highe
ac i i y in summe is asc ibed mos o en o high ai empe a u es and o
bi ing and pa asi ic insec s (Thing & Thing 1983, Mö schel & Klein 1997,
Coleman e al. 2000) and i has been documen ed ha wind speed a ec s
mosqui oes, which ha e a h eshold o 4-6 m/s (Russell e al. 1993, Pa o
2007). Inc eased ca ibou mo emen esul ing om insec ha assmen educes
o aging ime and nega i ely impac s cal g ow h and cow body condi ion
(Mö schel & Klein 1997, Cou u ie e al. 2009). Thus insec elie habi a , i.e.,
windy loca ions and snow pa ches (Joly & Klein 2011, Wilson e al. 2012) is an
impo an conside a ion when managing essen ial habi a s.
Following pa u i ion, i is ypical among Ce ids ha he e occu s an
immedia e and ma ked decline in daily mo emen (Long e al. 2009). The
sudden cessa ion o mo emen a bi hing by ca ibou cows was i s desc ibed
by Len (1966), who obse ed ha cows always s opped s ill when labou
commenced, ega dless i ha cow had been wi h a mo ing g oup, which le
he bi hing cow behind. Fe guson & Elkie (2004) obse ed he same
phenomenon, which was also la e suppo by Nagy’s (2011) mo emen a es
o cows a ound he ime o cal ing and known o la e ha e cal es-a -heel
(n=336). S ill, an assump ion ha a cow being s a iona y equa es wi h bi hing
equi es u he subs an ia ion. Video colla s, eco ding daily, on pa u ien
cows would be one solu ion.
Annual
The e was a dis inc seasonal dis ibu ion o he colla ed ca ibou cows in he
Cen al egion. Gene ally, he e was a sou hwes -no heas axis o hei
annual mo emen s. Al hough mo emen s we e o en sho and no mally no
om seacoas o Ice Cap, each AM cow was ypically u hes eas in summe
and u hes wes in win e . Also, each cow u ilized only a ac ion o he
a ailable a ea. This suppo s Cuyle & Linnell’s (2004) idea ha he e is a high
deg ee o popula ion sub-di ision o he AM ca ibou in he Cen al egion.
This knowledge is ele an o ca ibou managemen , e.g., conse a ion ac ions
51
applied o only a po ion o he egion will p obably no bene i he en i e
popula ion. Fu he , s ochas ic wea he e en s, which ha e he po en ial o
in luencing ca ibou abundance nega i ely, i limi ed in ex en o jus a po ion
o he egion, will also no likely a ec he en i e AM popula ion.
Mig a o y pa h leng h
AM cow mo emen s be ween win e and summe anges we e on a ela i ely
small scale. The mean ex en , 76 km, o AM cows’ mig a o y pa h leng h was
simila o ha obse ed in he 1990’s (Cuyle & Linnell 2004). In con as ,
ba en-g ound ca ibou (same sub-species as AM cows), and hose o no he n
Alaska, ypically make long mo emen s be ween win e and cal ing home
anges. These dis ances can be in he o de o 300-500 km o he Cen al
Alaskan o Po cupine he ds (C aighead & C aighead 1987, Fancy e al. 1989)
o many o he o he he ds in no he n Canada (Hall 1989). On a geog aphic
scale, he AM s udy a ea is much smalle han mos No h Ame ican
mig a o y ba en-g ound ca ibou home anges. Thus, ou obse ed sho AM
mig a o y pa hs we e no unexpec ed, and hey illus a e he con ined a eas
a ailable o ca ibou in Wes G eenland. Also e lec ed is he di e si y o
habi a s a ailable o he AM ca ibou ac oss ela i ely sho dis ances. Since
hey do no a e se la ge dis ances, AM cows could pa i ion any annual
ene gy budge su plus o hings o he han locomo ion, e.g., ep oduc ion.
This was suppo ed in he 1990’s when AM cows exhibi ed an excep ional
li e ime ecundi y (Cuyle & Øs e gaa d 2005).
Mo emen o colla ed AM cows be ween win e and summe anges was less
han 100 km, ollowed an ele a ion g adien , and hey we e spa ially
independen om each o he . Thus, AM cows appea unlike hei ba en-
g ound cousins and mos simila o he Dolphin & Union Island, No h
Ame ican Bo eal and Moun ain Woodland Ca ibou R. . ca ibou (Oosenbu g &
Thebe ge 1980, Be ge ud e al. 1984, Skogland 1986, 1989, Edmonds 1988,
Hillis e al. 1998, Nagy 2011, Nagy e al. 2011). This also highligh s he
impo ance o eleme y s udies.
Seasonal ac i i y pe iods
Indi idual ca ibou selec habi a o hei own success ul su i al and
ep oduc ion (Bélange & Cô é 2016, Fancy & Whi en 1991, Albon &
Lang a n 1992). Pa e ns o mo emen e ealed 10 seasonal ac i i y pe iods
o he G eenland AM popula ion. These we e associa ed wi h speci ic
52
habi a / ange a ibu es. Ou classi ica ion alls be ween Russell e al. (1993)
who iden i ied 15 ac i i y pe iods o mig a o y ba en-g ound ca ibou, and
Nagy’s (2011) eigh pe iods o bo h bo eal ca ibou and und a-win e ing
ba en-g ound ca ibou. Fu u e analyses may educe he pa i ioning o ewe
han 10 pe iods, since o example he habi a s chosen o ea ly, mid and la e
win e pe iods appea simila .
Speci ic ele a ion use was associa ed wi h he di e en seasons h oughou
he yea . Ele a ions unde 200 m we e always used o b eeding h ough o
he end o la e win e , while ele a ions abo e 350 m we e in a iably used o
cal ing and o he end o mid-summe . Ele a ion appea ed o be a p ima y
a ibu e de ining seasonal anges. This knowledge could be used o
conse a ion ac ions, e.g., ending an au umn ha es be o e he onse o he
b eeding season, when ca ibou a e a low ele a ions and accessible o hun e s.
La e summe and all anges p o ide he basis o a ca ibou’s body condi ion
build-up ha will pe mi pa icipa ion in he u (Came on e al. 1993), while
win e ange is i al o su i al (Be ge ud e al. 2007). Thus hese anges
quali y among he essen ial anges o ca ibou, i.e., impo an o
ep oduc ion o su i al. Un o una ely, o AM ca ibou, he e is a sca ci y o
habi a a ea associa ed wi h la e summe , all and win e ela i e o he o he
pe iods. Fo example, p e e ed AM win e habi a use is a ele a ions unde
200 m, and while win e ange is i al o ca ibou su i al, his habi a is in
sho supply o AM ca ibou. Win e ange is he smalles a ea o all he
seasonal anges. AM win e ange may he e o e need special a en ion and
p o ec ions. Inc easing he conce n, is ha he la ges ac o win e habi a ,
he Akia-No dland lowlands no h o God håbs jo d, is exposed o
sou hwes e ly s o m sys ems. These b ing he possibili y o se e al nega i e
wea he e en s, e.g., deep snows, ain-on-snow, and icing. The sho age o
la e summe , all and win e anges, indica es ha hese could be limi ing o
he AM ca ibou popula ion.
Seasonal habi a selec ion
Habi a selec ion a ied by season, and seasonal habi a s a ied conside ably
in hei dis ibu ion and a ea, ela i e o o al s udy a ea. Cal ing habi a had
he g ea es a ea and dis ibu ion. Tied o he leas a ea we e all/p e-
b eeding and la e all/ea ly win e . In ac , o he ac i i y pe iods om all
h ough mid-win e , habi a s we e ela i ely ew and small in a ea. The
53
la ges was he Akia-No dland lowlands, ollowed by he smalle lowlands o
he Na ssa ssuaq and Iluliak alleys. The e we e also a ew sca e ed smalle
alley lowlands. Taken oge he , hey comp ised only 21-26% o he o al
s udy a ea. The sca ci y o hese habi a s emphasizes ha human in luences
causing loss o hese could ha e de imen al impac s on he AM popula ion.
In e es ingly, he all/p e-b eeding ac i i y pe iod e idenced he lowes
pe cen age o cow loca ions e en in he highes occu ence ca ego y. An
ea lie s udy obse ed ha he lowes p obabili y o occu ence o cows was
a loca ions ha ing he highes alues o hun ing (Simonsen 2011). Since
all/p e-b eeding coincides wi h he hun ing season (1 Aug – 30 Sep ), esul s
may e lec a oidance o dis u bance / p eda ion isk om hun e s using he
same a eas. F om a managemen pe spec i e, his sugges s ha he human
dis u bance associa ed wi h a long summe /au umn ha es season may
nega i ely in luence b eeding and subsequen ly cal p oduc ion.
Ou analyses o p obabili y o cow occu ence du ing cal ing and b eeding
a e suppo ed by simila esul s ob ained by Simonsen (2011). Ou s udy,
howe e , was able o iden i y mo e a eas. As ega ds a compa ison o
summe and all habi a s, ou p obabili y o occu ence esul s sugges ha
all [Sep embe ] habi a / ange is limi ed in a ea/a ailabili y, while summe
[July] habi a / ange is no . The small bu impo an all a ea sugges s ha all
habi a may equi e p o ec ion measu es. This would be in addi ion o July
habi a , which is impo an o among o he hings selec i e eeding by cows
and insec elie (Tams o e al. 2005, Wilson e al. 2012). In he ollowing
sec ions, we will explo e linkages among o example nu i ion, o age, insec
elie , ele a ion, hun ing and win e snow.
Cal ing
Mean ele a ion use, ca. 400 m, in he cal ing ac i i y pe iod (11 May – 17
June) was signi ican ly highe han o all o he ac i i y pe iods. Cuyle &
Linnell (2004) and Simonsen (2011) also obse ed highes ele a ion use by
AM cows du ing cal ing. S ikingly, cal ing si es hemsel es we e ypically a
leas 200 m abo e and beyond he mean ele a ion use o all cows in he
cal ing ac i i y pe iod. This sugges s ha wha e e bene i s ele a ion con e s,
pa u ien cows appea o be maximizing hese. Cal ing si es also had a
gene ally sou he ly aspec , which would ake ad an age o sola adia ion.
54
Wi h he ad en o eliable aspec models, u u e s udies may be able o
explo e he signi icance o aspec on habi a selec ion a he landscape scale.
Simila o obse a ions in he la e 1990’s (Cuyle & Linnell 2004), AM cal ing
si es we e dispe sed in a con inuum ac oss he Cen al egion, whe e he
s aigh -line dis ance be ween seacoas and G eenland Ice Cap is only abou
110 km. Ce ainly 1/3 o bi hs occu ed wi hin 25 km o he G eenland Ice
Cap, howe e , se e al we e wi hin 10-20 km o he seacoas . Only 4-5 si es
co esponded wi h he ‘cal ing g ound’ delinea ed in NunaGis (2015).
Conside ing only inland a eas, as sui able o cal ing, appea s o ha e li le
ele ance o AM cow habi a selec ion a cal ing. Cow dispe sal a cal ing is
ypical o he mon ane eco ype o ca ibou (Be ge ud e al. 1984, Skogland
1989) and once again, highligh s how e y unlike he AM ca ibou a e om
hei ba en-g ound cousins. In con as o p oximi y o he Ice Cap, ele a ion
was he good p edic o o p obabili y o AM cow occu ence a cal ing (and
b eeding).
The AM cal ing con inuum is unsu p ising gi en ha 42% o his ange
possesses he habi a a ibu es associa ed wi h highes p obabili y o
occu ence by cows du ing peak cal ing: high ele a ion, hea h, open hea h,
copse, en, g ass, snowbed and wind exposed idge. Po en ial xe ic habi a o
cal ing is p esen nea he Ice Cap, bu he ex en o essen ial cal ing habi a
a ailable elsewhe e h oughou he egion is now ecognized. This
s eng hens capaci y o egula e human ac i i ies in cal ing habi a be o e
du ing and a e cal ing.
Based on daily mo emen a es, he seasonal ac i i y pe iod o cal ing las ed
om 11 May un il 17 June. Using ac ual bi hing e en s, ‘Peak’ cal ing (68%)
was a 19-day pe iod (21 May-8 June), wi h ‘mos ’ cal ing (95%) o e a 32-day
pe iod, 14 May o 15 June. The pe iod leng h o AM peak cal ing is simila o
Bo eal ca ibou (R. . ca ibou) (Nagy 2011). May bi hs we e unexpec edly
common, mean 30 May ± 8.2 days. Gene ic mixing wi h e al semi-domes ic
eindee (Jepsen e al. 2002), which cal in May, is a plausible explana ion o
he ea ly cal ing. Changing wea he o plan phenology migh also be ac o s.
We no e ha mean cal ing da e 2010 was 7-10 days ea lie han 2008,
howe e , he small 2010 sample size p ecludes conclusions abou a o wa d
shi in cal ing phenology.
55
Cal ing si e ideli y
High ideli y o cal ing g ounds illus a es hei impo an ole in he annual
cycle o cal p oduc ion (Gunn & Mille 1986, Fancy & Whi en 1991, Schae e
e al. 2000, Mahoney & Schae e 2002, Russell e al. 2002, Fe guson & Elkie
2004). The sa elli e-colla ed AM cows exhibi ed high ideli y o p e iously
chosen cal ing si es. Howe e , ou small sample size and sho s udy pe iod
do no pe mi conclusi e s a emen s ega ding pa e ns. I is in e es ing o
no e ha ou minimum alue o 0.5 km and median alue o 7.1 km we e
simila o he 4 o 11 km obse ed o ca ibou o he moun ain eco ype in
sou heas e n Lab ado (Popp e al. 2011) and he < 5 km o woodland ca ibou
o no he n On a io (Fe guson & Elkie 2004). In con as , Yukon-Alaskan
Po cupine ca ibou cows, which hough ai h ul o a gene al a ea did no cal
nea same loca ion annually, e.g., minimum 67.1 ± 49.1 km (Fancy & Whi en
1991). Gi en he high ideli y shown by AM cows, an h opogenic dis u bance
could ha e s ong nega i e impac s i cows a e o ced o educe ideli y o
p e e ed si es (Faille e al. 2010).
Cal ing si e choice
Sp ing mig a ion o pa u ien Rangi e cows is o en no hwa d in la i ude, o
owa ds inc easing ele a ions. This is hough o be an e olu iona y esponse
o empo al and spa ial esou ce a ia ion ha will maximize cal su i al,
e.g., inc ease a ailable o age quan i y and quali y, o educe he isk o
neona e p eda ion (Be ge ud & Page 1987, F yxell e al. 1988, Klein 1990,
Albon & Lang a n 1992, G i i h e al. 2002, Russell e al. 2002, Pos e al. 2003,
Loe e al. 2005). The e is no doub ha cal ing a eas a e essen ial o
p oduc i i y o a ca ibou popula ion, and ha access o hese has di ec
consequences a he popula ion le el (Ca oll e al. 2005).
Ca oll e al. (2005) ound ha cal ing si es we e p ima ily ela ed o he
iming and p esence o snowmel . Pa u i ion and subsequen lac a ion
ep esen eno mous ene gy expendi u es o cows (Boe je 1985, Be ge ud e
al. 2007), ye pa u ien ca ibou employ a capi al ep oduc i e s a egy, i.e.,
ni ogen demands h oughou ges a ion and ea ly lac a ion a e chie ly
co e ed by ma e nal body ese es, which we e deposi ed la e in he p e ious
g owing season (Moen e al. 2006, Ba boza & Pa ke 2008, Taillon e al. 2013).
Thus, peak pa u i ion can p ecede g een-up by se e al weeks (Du an e al.
2005, Moen e al. 2006, Ba boza & Pa ke 2008, Taillon e al. 2013), and in ac ,
pa u ien cows no mally a i e a cal ing a eas be o e g een-up has occu ed
(Whi en & Came on 1980, Kellyhouse 2001). Once g een-up begins, cal ing
56
a eas a e associa ed wi h high a es o biomass inc ease (Kellyhouse 2001),
and he eme gen plan s a e high in soluble ca bohyd a es, ni ogen and
phospho us, which de e io a e as summe ad ances (Whi e e al. 1975,
Whi en & Came on 1980, Jo gensen e al. 2002). Thus, eme gen o age a
cal ing si es may well coincide wi h he bi hing cows’ pe iod o g ea es
ene gy needs, which is abou h ee weeks pos -cal ing (Pa ke e al. 1990).
As ega ds a ailable o age quan i y and quali y a cal ing, ugged e ain,
e en in lowlands, widens he pe iod o eme gen o age a ailabili y in a eas
o pa chy snowmel (Nelleman & Thomsen 1994). Thus, high ele a ion would
no be a p e equisi e o eme gen o age on he ough- ea u ed AM egion.
AM lowlands can ha e pa chy snowmel wi h eme gen ege a ion
unde way, ye mos pa u ien adul AM cows bi hed a ela i ely high
ele a ions o ca. 600 m o mo e. Gi en mean cal ing da e, 30 May, and he ca.
65°N la i ude o he Cen al egion, eme gen ege a ion is unlikely a he
ele a ions ypically chosen o bi hing by AM cows. AM cal ing appea s o
occu whe e ood esou ces a e limi ed and ege a ion g een-up many weeks
in he wai ing, e.g., maximum NDVI’s (no malized di e ence ege a ion
index, ‘densi y o g een’) ac oss he Cen al egion gene ally do no occu
un il July (Tø up 2009).
Why would AM cows choose high ele a ions o xe ic a eas? Why no bi h in
he snowmel occu ing in he Cen al egion’s lowlands? The la e would
pe mi access o eme gen o age, which as sp ing p og essed could be
pu sued up he eadily a ailable ele a ion g adien s h oughou he egion.
Since insec ha assmen i s begins se e al weeks pos -cal ing, escape om
hese is no likely a ac o in bi hing si e choice. Insec elie could, howe e ,
keep cows a high ele a ions, which a e o en windy, in he pos -cal ing
pe iod. Dec easing he neona e p eda ion isk is a mo e poin ; because la ge
p eda o s ha e been absen in Wes G eenland since a leas he 1870’s (Vibe
1981). The ele a ions chosen by AM cows sugges he possibili y o g ea e
nu ien alue con ained in eme gen ege a ion associa ed wi h snowmel in
highlands, and ha i is a ailable o e a longe pe iod. Fu u e in es iga ion o
he nu ien alue and pe iod leng h o eme gen ege a ion a high ele a ion
e sus lowlands may cas ligh on his conund um.
Wea he migh also in luence cal ing si e choice. The Akia-Manii soq egion
is exposed o sou h wes e ly s o m sys ems. Bi hing cows choosing high
ele a ions o d y xe ic a eas close o he G eenland Ice Cap may be educing
57
he isk o un a ou able wea he condi ions o neona e su i al, e.g., ain,
which could inc ease neona e mo ali y h ough excessi e hea loss associa ed
wi h cold s ess hypo he mia. Hea y ain all and wind can d as ically
inc ease neona e mo ali y in A c ic animal popula ions (Blix & S een 1979,
Mallo y e al. 2009, Anc il e al. 2014, Yannic e al. 2014) including ca ibou
neona es (Kelsall 1968, Blix 1980). E en wi hou wind, we neona e hea loss
inc eases 5- old (Len z & Ha 1960, Ha e al. 1961, Ma kussen e al. 1985).
Al hough ain coupled wi h wind inc eases neona e mo ali y, i s ole on
pa u ien ca ibou mig a ion and cal ing si e choice has been igno ed (Mille
& Gunn 1986). Reducing he likelihood o exposing neona es o ain and we
subs a es would bene i o sp ing su i al in la ge ungula es (Azzam e al.
1993). This hypo hesis ha high ele a ion (combined wi h 65°N la .) o xe ic
a eas p omo es neona e su i al by educing he isk o ain, needs u he
s udy. I ue, his may cla i y wha op ions a e open o pa u ien cows on
anges wi h low elie landscapes whe e he only ain a oidance choices may
be loca ions o d ies clima e o he mos no he ly la i udes possible.
In e es ingly, ha ing a yea ling-a -heel (cal om he p e ious sp ing)
appea ed o a ec a cow’s ele a ion choice (median 186 m lowe ) o
pa u i ion. The di e ence app oached signi icance (P = 0.058, d = 10, =
2.2281) and may be biologically signi ican . While cows a e capi al b eede s,
i.e., ely on body ese es o bi hing and ini ial lac a ion, he yea ling-a -heel
may ha e ewe body ese es emaining a win e ’s end, making high
ele a ions, snow-bound, wi h educed o age a ailabili y less han ideal.
Pa u ien cows wi h yea ling-a -heel om he p e ious sp ing may choose
bi hing si es a lowe ele a ions o balance hei yea ling’s o age needs
agains enhancing su i al o he coming neona e. To da e, he e a e no
s udies add essing pa u ien ca ibou cal ing si e choice in he p esence o
absence o a yea ling-a -heel. Ae ial obse a ion o pa u ien cows p e-
cal ing combined wi h in es iga ions o p e- and pos -cal ing eloca ions,
migh asce ain i his ade-o is occu ing.
Managemen Implica ions
Mo ali y 2008-2010
Adul emale and cal su i al a e he wo key demog aphic ac o s ha
de e mine popula ion g ow h a es (Ha e & Be ge ud 1991). The 2008-2010
mo ali y among he AM sa elli e-colla ed cows was dis u bingly high: 26 o
64
penal ies o illegal ha es and he inancial esou ces was ed when a
colla ed cow is emo ed un imely om eleme y s udies. Hun e ha es
exace ba ed colla ed-cow mo ali y in he cu en s udy. An in o ma ion
campaign could inc ease public unde s anding o he signi icance o colla
da a o e ec i e managemen o habi a and ca ibou o he bene i o all.
Acknowledgemen s
The A c ic En i onmen P og am o he Danish Minis y o En i onmen and
Ene gy, he aluminium smel e company ALCOA, he Danish Assis ance o
he A c ic P og am (DANCEA), he G eenland Go e nmen and he
G eenland Ins i u e o Na u al Resou ces p o ided unding. The o me we e
ob ained by Pe e Aas up. Sa elli e-based ege a ion maps we e p o ided by
ALCOA. The G eenland go e nmen ’s Asiaq aided us wi h he digi al
ele a ion model. We hank Ai G eenland Cha e and hei pilo s o sa e
lying du ing ca ibou cap u e. We hank Ba y Mino o e icien ne -gunning
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B ooke ad ised on unsui abili y o linking passi e mic owa e a iables o
poin da a. We hank Wendy Loya o ex ensi e e iew o he manusc ip and
Joseph McCullough o p oo eading.
65
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God håbs jo den. Geog aphic Resou ce Analysis and Science L d, Ins i u e o Geog aphy,
Uni e si y o Copenhagen Technical epo o NERI/Alcoa. 27pp. www.g as.ku.dk
80
Table 19. Pos -cal ing ca ibou esou ce selec ion models
Ten bes - i pos -cal ing ac i i y pe iod esou ce selec ion models
SBIC
∆SBIC
Rela i e
p e e ence o
bes - i model
based on
∆SBIC*
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-136853.8
0
Bes - i model
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-136842.2
12
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock
-136783.2
71
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen
-136780.9
73
Ve y s ong
hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-136779.9
74
Ve y s ong
hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-136778.0
76
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed
-136775.6
78
Ve y s ong
hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-136770.6
83
Ve y s ong
hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-136768.5
85
Ve y s ong
ele a ion + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-136766.0
88
Ve y s ong
*Rela i e p e e ence o bes - i model based on Ha din & Hilbe (2012).
81
Table 20. Ea ly summe ca ibou esou ce selec ion models
Ten bes - i ea ly summe ac i i y pe iod esou ce selec ion models
SBIC
∆SBIC
Rela i e
p e e ence o
bes - i model
based on
∆SBIC*
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-140048.1
0
Bes - i model
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-140036.7
11
Ve y s ong
ele a ion + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-140012.2
36
Ve y s ong
ele a ion + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-140005.7
42
Ve y s ong
ele a ion + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-140000.1
48
Ve y s ong
hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-139994.3
54
Ve y s ong
ele a ion + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-139992.6
56
Ve y s ong
hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-139992.2
56
Ve y s ong
copse + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-139987.6
61
Ve y s ong
hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-139985.1
63
Ve y s ong
*Rela i e p e e ence o bes - i model based on Ha din & Hilbe (2012).
82
Table 21. Mid /la e summe ca ibou esou ce selec ion models
Ten bes - i mid/la e summe ac i i y pe iod esou ce selec ion models
SBIC
∆SBIC
Rela i e
p e e ence o
bes - i model
based on
∆SBIC*
ele a ion + hea h + open + hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-140439.1
0
Bes - i model
hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-140410.6
29
Ve y s ong
ele a ion + hea h + open + hea h + copse
-140408.5
31
Ve y s ong
ele a ion + hea h + open + hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-140406.6
33
Ve y s ong
ele a ion + hea h + open + hea h + copse + en + g ass
-140406.2
33
Ve y s ong
hea h + open + hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-140402.4
37
Ve y s ong
ele a ion + hea h + open + hea h + copse + en
-140401.9
37
Ve y s ong
ele a ion + hea h + open + hea h + copse + en + g ass + snow-bed
-140398.6
41
Ve y s ong
ele a ion + hea h + open + hea h + copse + en + g ass + snow-bed + wind-exposed
-140398.0
41
Ve y s ong
hea h + copse + en + g ass
-140391.4
48
Ve y s ong
*Rela i e p e e ence o bes - i model based on Ha din & Hilbe (2012).
83
Table 22. Fall /p e-b eeding ca ibou esou ce selec ion models
Ten bes - i all/p e-b eeding ac i i y pe iod esou ce selec ion models
SBIC
∆SBIC
Rela i e
p e e ence o
bes - i model
based on
∆SBIC*
hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-137204.7
0
Bes - i model
hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-137198.2
7
S ong
hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-137196.2
9
S ong
hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-137190.5
14
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-137188.8
16
Ve y s ong
hea h + copse + en + g ass
-137188.4
16
Ve y s ong
hea h + open-hea h + copse + en + g ass
-137187.4
17
Ve y s ong
hea h + copse + en + g ass + snow-bed
-137184.4
20
Ve y s ong
hea h + open-hea h + copse + en + g ass + snow-bed
-137183.2
22
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-137180.9
24
Ve y s ong
*Rela i e p e e ence o bes - i model based on Ha din & Hilbe (2012).
84
Table 23. B eeding ( u ) ca ibou esou ce selec ion models
Ten bes - i b eeding ac i i y pe iod esou ce selec ion models
SBIC
∆SBIC
Rela i e
p e e ence o
bes - i model
based on
∆SBIC*
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-143218.3
0
Bes - i model
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-143213.1
5
Posi i e
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed
-143126.4
92
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock
-143118.2
100
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen
-143117.2
101
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed
-143111.7
107
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass
-143093.9
124
Ve y s ong
hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-143043.9
174
Ve y s ong
hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-143034.5
184
Ve y s ong
ele a ion + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-143034.0
184
Ve y s ong
*Rela i e p e e ence o bes - i model based on Ha din & Hilbe (2012).
85
Table 24. Pos -b eeding /la e all ca ibou esou ce selec ion models
Ten bes - i pos -b eeding/la e all ac i i y pe iod esou ce selec ion models
SBIC
∆SBIC
Rela i e p e e ence o
bes - i model based on
∆SBIC*
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-154203.3
0
Bes - i model
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-154196.2
7
S ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen
-154135.9
67
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock
-154128.7
75
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed
-154081.3
122
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed
-154079.6
124
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass
-154048.3
155
Ve y s ong
ele a ion + hea h + open-hea h + copse + en
-153983.9
219
Ve y s ong
ele a ion + hea h + open-hea h + copse
-153957.5
246
Ve y s ong
ele a ion + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-153946.3
257
Ve y s ong
*Rela i e p e e ence o bes - i model based on Ha din & Hilbe (2012).
86
Table 25. Ea ly/mid-win e ac i i y pe iod ca ibou esou ce selec ion models
Ten bes - i ea ly/mid-win e ac i i y pe iod esou ce selec ion models
SBIC
∆SBIC
Rela i e
p e e ence o
bes - i model
based on
∆SBIC*
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-144092.0
0
Bes - i model
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-144085.8
6
Posi i e
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock
-144036.5
56
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen
-144031.9
60
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed
-144021.2
71
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed
-143994.7
97
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass
-143983.1
109
Ve y s ong
ele a ion + hea h + open-hea h + copse
-143973.9
118
Ve y s ong
ele a ion + hea h + open-hea h + copse + en
-143971.0
121
Ve y s ong
ele a ion + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-143962.3
130
Ve y s ong
*Rela i e p e e ence o bes - i model based on Ha din & Hilbe (2012).
87
Table 26. La e win e ac i i y pe iod ca ibou esou ce selec ion models
Ten bes - i la e win e ac i i y pe iod esou ce selec ion models
SBIC
∆SBIC
Rela i e
p e e ence o
bes - i model
based on
∆SBIC*
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-139452.9
0
Bes - i model
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-139451.2
2
Weak
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed
-139417.2
36
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock
-139413.4
40
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen
-139411.0
42
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed
-139402.4
51
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass
-139387.0
66
Ve y s ong
ele a ion + hea h + open-hea h + copse + en
-139377.5
75
Ve y s ong
ele a ion + hea h + open-hea h + copse
-139370.1
83
Ve y s ong
ele a ion + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-139345.7
107
Ve y s ong
*Rela i e p e e ence o bes - i model based on Ha din & Hilbe (2012).
88
Table 27. P e-cal ing ac i i y pe iod ca ibou esou ce selec ion models
Ten bes - i p e-cal ing ac i i y pe iod esou ce selec ion models
SBIC
∆SBIC
Rela i e
p e e ence o
bes - i model
based on
∆SBIC*
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-137443.7
0
Bes - i model
ele a ion + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e
-137442.2
2
Weak
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-137440.7
3
Posi i e
ele a ion + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-137439.6
4
Posi i e
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock
-137431.4
12
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen
-137425.4
18
Ve y s ong
ele a ion + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock
-137424.8
19
Ve y s ong
ele a ion + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen
-137418.6
25
Ve y s ong
ele a ion + hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed
-137406.5
37
Ve y s ong
hea h + open-hea h + copse + en + g ass + snow-bed + wind-exposed + soil/ ock + sedimen + wa e + snow/ice
-137406.3
37
Ve y s ong
*Rela i e p e e ence o bes - i model based on Ha din & Hilbe (2012).
89
Appendix 3
Habi a selec ion by G eenland ca ibou (Rangi e a andus g oenlandicus) du ing he CALVING pe iod ela i e o hose
selec ed du ing he o he ac i i y pe iods, based on la en selec ion di e ence (LSD) unc ion model compa isons, 2008-2010.
Table 28. Cal ing VERSUS pos -cal ing, ea ly summe , and mid /la e summe ac i i y pe iods
Habi a ypes
Cal ing e sus pos -cal ing
Cal ing e sus ea ly summe
Cal ing e sus mid /la e summe
Coe icien
S.E.
P- alue
RS*
Coe icien
S.E.
P- alue
RS
Coe icien
S.E.
P- alue
RS
hea h
-0.301
0.094
0.001
26
0.029
0.084
0.732
=
-0.661
0.094
0.000
48
openhea h
0.520
0.098
0.000
>100
0.858
0.084
0.000
>100
0.478
0.085
0.000
>100
copse
0.059
0.103
0.567
=
0.112
0.094
0.233
=
-0.004
0.085
0.964
=
en
-0.130
0.100
0.192
=
-0.200
0.087
0.022
18
-0.093
0.090
0.302
=
g ass
0.120
0.076
0.113
=
0.145
0.070
0.039
>100
0.134
0.071
0.059
=
snowbed
-0.366
0.170
0.032
31
-0.429
0.143
0.003
=
-0.488
0.141
0.001
39
windexp
-0.003
0.097
0.974
=
0.378
0.087
0.000
>100
0.842
0.085
0.000
>100
soil ock
0.042
0.084
0.616
=
-0.102
0.077
0.188
=
-0.355
0.081
0.000
30
sedimen
-0.170
0.121
0.158
=
-0.372
0.099
0.000
31
-0.328
0.103
0.001
28
wa e
0.310
0.139
0.026
>100
0.065
0.115
0.572
=
-0.092
0.116
0.428
=
snowice
0.889
0.784
0.257
=
0.297
0.466
0.524
=
-
-
-
-
_cons
0.700
0.133
0.000
-0.230
0.117
0.050
0.483
0.121
0.000
*Rela i e selec ion (RS) was calcula ed o a iables wi h coe icien s signi ican ly di e en om 0 as exp (b) when b>0 and as [1-exp(b) when b<0 (La ham, La ham &
Boyce, 2011b). Rela i e selec ion alues <100 indica e ha use o he habi a du ing cal ing was signi ican ly less by x% han ha o he season being compa ed,
alues >100 indica e use o habi a du ing cal ing was signi ican ly g ea e han ha o he season being compa ed, and = indica es habi a use du ing cal ing and
season being compa ed we e no signi ican ly di e en .