65
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Field a icle
Shi happens – a glimpse in o males’ ma ing ac ics in a polygynous
ungula e - he eindee
Øys ein Holand1, Léon L’I alien2, Robe B. Weladji3, Na asa Djako ić1, Knu Røed4, Anne
Lene Ho land1, & Mau i Nieminen5
1 Depa men o Animal and Aquacul u al Sciences, No wegian Uni e si y o Li e Science, P.O.Box 5003, N-1432, Ås,
No way (co esponding au ho : [email p o ec ed]).
2 Dépa emen de Biologie and Cen e d’é udes no diques, Uni e si é La al, 1045 a enue de la Médecine, Québec,
Québec, G1V 0A6, Canada.
3 Depa men o Biology, Conco dia Uni e si y, 7141 She b ooke S . Wes , Mon eal, QC, H4B 1R6, Canada.
4 Depa men o Basic Sciences and Aqua ic Medicine, No wegian School o Ve e ina y Science, P.O.Box 8146, Dep.
N-0033 Oslo, No way.
5 Finnish Game and Fishe ies Resea ch Ins i u e, Reindee Resea ch S a ion, Fin-99910 Kaamanen, Finland.
Abs ac : This is abou he wo big “guys”, Spo and Mika, and hei endea ou o p opaga e hei genes du ing he ma -
ing season 2007. They we e 6 and 5 yea s old weigh ing 172 and 141 kg be o e u , espec i ely. Toge he wi h 23 o he
males (one 5 y s, wo 4 y s, h ee 3 y s, six 2 y s and ele en 1 y old) hey oamed wi hin a ~15 km2 enced a ea compe ing
o access o 87 emales. Indeed, he compe i ion was in ense and all males p esen con ibu ed o he dynamic obse ed.
Especially Ho , he hea ies 4 y s old male weighing 155 kg be o e u , played a p ominen ole – in addi ion o Spo and
Mika, hei ma ing ac ics being highly dynamic. Howe e he e is no sho cu o success – s eng h ha e o be coupled
wi h sma ness - bu shi happens - as we’ll see.
Key wo ds: in asexual selec ion; male-male comba s; ma ing ac ics; eindee ; eindee male ma ing s a egy, ideos.
Rangi e , 32 (1): 65–72
Ri al y
P elude - Spo ’s ea ly ise
Spo s a ed u ing ea ly (Pho o 1). Al eady in mid
Sep embe he s a ed o show o ; he ding emales
as well as chasing i als while g un ing con iden ly.
Towa ds he end o Sep embe his u ing ac i i y
s eadily in ensi ied (Fig.1a), con olling a g owing
ma ing g oup (Fig. 1c). This coincided wi h his i s
ma ing a emp (Fig. 2), sugges ing ha he emales
we e s a ing o come in o hea . Con olling such a
big emale g oup always on he mo e sea ching o
ood, while 10-15 o he males we e hanging a ound
wai ing o hei chance, ook all his a en ion.
Indeed, Spo aded eeding wi h keeping con ol as
con i med by he s ong in e se ela ionship be ween
ale s anding and o aging.
Mika (Pho o 2) showed a di e en p elude. His
ole ance owa ds o he males and limi ed he ding
beha iou kep spending low and le mo e ime
a ailable o o aging (Fig. 1b). As a consequence,
NOR’s 16 h No dic Con e ence on Reindee and Reindee Husband y Resea ch
T omsø, No way, 16 h-18 h No embe 2010
66 Rangi e , 32 (1), 2012
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his p e u ma ing g oup was small as compa ed o
Spo ’s (Fig. 1c). Mos o he o he p ime- (4 & 5 y s)
and semi p ime-aged (3 y s) males we e also ac i ely
ying o o m ma ing g oups, as opposed o he
youngs e s (1 and 2 y s old males), bu hei g oups
we e labile.
The hea is on
Spo ’s in ole ance owa ds i als climaxed Oc o-
be 2nd as he he d en e ed he peak u (2nd – 9 h
Oc obe ). Especially he inexpe ienced 1 y old males
we e igo ously chased by Spo as hey eage ly and
op imis ically app oached “his” emales. Mika who
joined Spo ’s g oup Oc obe 1s , kep a low p o ile
(Fig. 1). He was wa ching he “ballgame” ca e ully,
bu ning as li le uel as possible and hoping o he
“jackpo ” – o ake o e he whole ma ing g oup now
consis ing o a ound 50 emales (Fig. 1c), o which
many we e s a ing coming in o oes us.
Spo was igilan ly s anding wa ching and g un -
ing (Fig. 1a) and a he same ime eage ly chasing
i als and he ding as well as inspec ing emales.
Howe e , his in ense ma ing beha iou c ea ed
unease among he emales. They sp ead ou o mi i-
ga e he annoyance. This se iously s e ched his
con olling capaci y. We could ac ually “ eel” ha
his body ese es we e sh inking. Indeed, his ac ic
paid o as con i med by se e al success ul copula-
ions obse ed he i s days o Oc obe (Fig. 2). Bu ,
would Spo be able o keep up his pace h ough he
peak u ? I no , who would en e he cen e cou ?
O would he big ma ing g oup s a o disin eg a e?
Ho en e s he scene - Mika - he “sma y” - akes con ol
Oc obe 3 d in he ea ly a e noon Ho suddenly
a i ed oge he wi h a small g oup o emales and
ook on Mika who was ope a ing in he ou ski o
Spo ’s g oup, igh away (click o ideo 1). Ho pu
up a ough igh bu inally ga e in and an away.
Howe e , i een minu es la e he e u ned - i o
a igh - and challenged Spo , he big boss (Pho o
3) (click o ideo 2). A e se en minu es igh ing
b a ely Ho had o h ow in he owel. Immedia ely,
Mika esh a e he wa ming up ound wi h Ho 20
minu es ea lie , challenged Spo , s ill punch-d unk,
and knocked him ou a e a sho ( wo minu es)
in ense igh (click o ideo 3).
In he e ening he dus had se led. Mika was in
con ol o he big ma ing g oup, now consis ing o 65
emales. In addi ion, 16 i als we e swa ming a ound.
Oc obe 4 h Spo was able o keep a small ma ing
g oup o a ound 15 emales, whe eas he es (~40)
s ayed wi h Mika (Fig. 1c). The nex day he wo
g oups had me ged again (Fig. 1c). Spo s ayed in he
ou ski . His collapse seemed comple e and his ac ic
shi ed acco dingly; hanging a ound o see wha hap-
pens. Ac ually, he s a ed o aging (Fig. 1b) and ied
o ge going. Ho also e u ned - s ill “ho ” -, bu was
e ec i ely con olled by Mika. He had lea ned his les-
son and s ayed in he pe iphe y o he g oup o he
es o he peak u .
Pho o 1. Spo wi h some ladies. Pho o: Na asa Djako ić.
Pho o 2. Mika. Pho o: Léon L’I alien.
Pho o 3. Spo & Ho igh ing. Pho o: Léon L’I alien.
67
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Mika’s heydays
Mika looked con iden ly subduing his i als, appa -
en ly con olling hem wi hou bu ning oo much
uel. The educed chasing beha iou , as compa ed o
Spo , seemed o calm down he emales in he g oup
and hey we e able o o age and es undis u bed.
Some o he young males ied o ake on emales in
he ou ski o he ma ing g oup bu hey seemed
o p e e o s ay wi h Mika and mo ed owa ds he
cen e when ha assed. This eased his he ding e o
as he could pa ol a a he condensed emale g oup.
Al hough he emales we e a he calm and Mika
ac ed con iden ly, i was impossible o him o de end
such a big mobile g oup om in uding males.
Hence, Mika displayed a mixed ac ic, ying o keep
he emales igh by being ole an owa ds i als
and he ding he emales gen ly and a he same ime
keeping a special eye on he emale wi hin he g oup
Fig. 1. Daily a e age; g un ing ac i i y based on 5 minu es con inuous obse a ions (a) and p opo ion spen eeding (b)
based on 15 minu es ocals wi h obse a ions e e y 15 seconds and emale ma ing g oup size ( he dominan male
is gi en he sco e o all emales wi hin he g oup) (c) o Spo and Mika du ing p e (g ay), peak (whi e) and pos
(g ay) u . Squa es ep esen dominance wi hin a ma ing g oup, iangles subdominance, whe eas ci cles deno e
Mika being se iously inju ed.
SPOT
p e peak pos
a)
b)
c)
Da e Da e
MIKA
p e peak pos
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closes o ecep i eness. He ac ually le o he p ime
and semi-p ime males ake pa in he “ o spiel”.
When he emale app oached ecep i eness Mika
in e e ed, ook o e he ending and cou shipping
and copula ed he . A e mission comple ed, he los
in e es and sough new emales coming in o hea .
Game o e o Mika
Oc obe 7 h we obse ed Mika limping. The day a e
we ound him se iously a ec ed (click o ideo 4).
He was no able o pu any weigh on his le on
leg. The inju y made him less mobile and cons ained
his abili y o exp ess his supe io i y, as e lec ed in
his dec eased u ing beha iou (Fig. 1a) and educed
ma ing g oup size (Fig. 1c). I was only a ma e o
ime be o e he would be o e h own, as also indi-
ca ed by he inc eased u ing beha iou o his i als.
Oc obe 9 h we ound Mika alone – es ing. The nex
day he was seen oge he wi h 5 emales (Fig. 1c), no
capable o ollowing hem as hey mo ed on, and he
was le behind.
Spo ’s Indian summe
Spo appea ed like a bi d Phoenix, as seen by his
inc eased g un ing ac i i y om 8 h o 9 h Oc obe
(Fig. 1a). He had ob iously used his “ ime ou ” e i-
cien ly, o aging (Fig. 1b) and es ing and had ecu-
pe a ed. Bu , his pace was uned down as he he d
was en e ing he pos u pe iod (Oc obe 10 h-15 h)
(Fig. 1a). Indeed, his seemed o be he case o all
he p ime and semi-p ime males. They had los mos
o hei ese es compa ed o he youngs e s, which
s ill we e in p e y good shape s ill eage ly checking
emales. The la es copula ion a emp obse ed was
Oc obe 10 h (Fig. 2). Ob iously he e was li le le o
igh o and he big ma ing g oup s a ed o disin-
eg a e (Fig. 1c). Indeed, males’ ocus shi ed and was
uned owa ds cha ging hei ba e ies (i.e. o aging)
(Fig. 1b) be o e he onse o win e .
Re lec ions
The winne doesn’ ake i all
Based on he numbe o success ul copula ions seen
and he ma ing g oup dynamic obse ed, Spo and
Mika ob iously seemed o be he wo mos success ul
males, as hey al e na ed o domina e he main g oup
consis ing o a ound 50 emales (Fig. 1c) du ing peak
u . Spo and Mika we e equally success ul, si ing
15 and 14 o sp ing, espec i ely (Fig. 3). Thus he
numbe o days o p imacy and he ma ing g oup
size du ing peak u e lec ed hei success, as also
epo ed in o he polygynous species (Pembe on e
al., 1992; Wade & Shus e , 2004). Ne e heless hey
e ilized only a ound 60% o he emales in he main
g oup, sugges ing ha some o he ~15 males hang-
ing a ound also go hei sha e.
Timing o he e o
The highly synch onised emale o ula ion in ein-
dee , abou 90% o he copula ion a emp s we e
obse ed Oc obe 2nd – 9 h (Fig. 2), induces s ong
selec ion p essu e o iming males’ ep oduc ion
e o , including hei igh ing abili y, wi h emales’
ep oduc i e phenology, as epo ed in o he no he n
ungula es (P es on e al., 2001; Mys e ud e al., 2008).
Indeed, all p ime- and semi p ime-aged males we e
a hei op du ing peak u , al hough some o hem
s a ed hei ma ing beha iou ea lie han o he s and
beha ed di e en ly. As eindee emales a e ecep i e
only one o wo days (Rops ad, 2000), he ope a ional
sex a io (OSR), de ined as he a io o e ile males
o ecep i e emales, du ing he peak u pe iod
(Oc obe 2nd – 9 h), will a e age abou i e imes he
absolu e sex a io. Gi en ha he emales dis ibu e
andomly be ween ma ing g oups, i espec i ely o
hei ep oduc i e phase, he local (wi hin g oup)
OSR will be equally skewed du ing peak u pe iod,
inducing in ense local male-male compe i ion.
A lexible s a egy
The males al e na ed be ween di e en ac ics based
on local spa ial and empo al a ia ion in OSR and
own s a us, as seen in many ungula es ( e iewed by
Is a an, 2005). Indeed, he emale oes us asynch ony
and he leng h o he ecep i eness “window” seem
su icien o keep up he en i onmen al po en ial
o polygyny (Emlen & O ing, 1977) in eindee . By
keeping he emales igh and de ending hem om
in ude s, he dominan male wi hin a ma ing g oups
was “p epping” he emales collec i ely. Ac ually also
Fig. 2. Da es o obse ed copula ion a emp s by Spo
(whi e), Mika (black) and o he males (g ay) in he
main ma ing g oup.
69
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his i als con ibu ed o he p e-cou ship and go
hei sha e. The dominan males ied o keep he
se icing ime low (no mally 2-4 h s du ing peak u )
o be eady o seek o new ecep i e emales wi hin
hei g oup asap. The emales, on he o he hand,
may be eage o ex end he se icing ime o eassu e
ha he dominan male is he igh guy. A longe
se icing ime will also educe o he emales access o
him and hence induce emale- emale compe i ion o
high quali y males. This may ac ually explain some
o he sp ead in ep oduc i e success among males
which a ilia ed wi h he main g oup du ing peak
u (Fig. 3). Indeed, ma e compe i ion and emale
choice do in e ac (B o-Jø gensen, 2011), as seen by
he emale in e es in Mika (click o ideo 4).
Sneaking – an age speci ic ac ic
Gi en he in ense local male-male compe i ion du -
ing peak u , Spo and Mikka ocused p ima ily on
con olling he mos po en i als, paying less a en-
ion o he youngs e s al hough hey we e o en a -
ge ed and chased as hey da ed o mo e a he eely
wi hin he ma ing g oup. This ac ually enabled mos
( ou ) o he 2 y s and some ( h ee) o he 1 y old
males s aying mainly in he main g oup du ing peak
u o sneak a ound and “bush” ma e a ew (one, wo
o h ee) emales each, adding up o 13 cal es o ally
(Fig. 3). A simila pa e n has also been epo ed by
Røed e al. (2002). Ob iously, o a he cal es a an
ea ly age gi es an ex a i ness bonus and induces age
speci ic selec i e p essu e o adop a sneaking ac ic
when young as hey we e no able o o m and con ol
own g oups in his highly compe i i e en i onmen .
Howe e , in en i onmen s wi h only young (1 y )
males a ailable hey may shi o a dominan ac ic,
al hough hei he ding abili y and display is no ully
de eloped (Holand e al., 2006).
Condi ional and condi ion dependen s a egy
Also he h ee semi p ime males (K9, K5 & K3) ope -
a ed mainly wi hin he main g oup and p ac ised an
al e na i e ac ic; s anding wa ching, ying o o m
empo al g oups o he d o single emales close o
oes us, si ing 14 cal es o ally (Fig. 3). Ho was he
big lose wi h no si e a all. He was wa ched in en-
si ely and aken “ou ” by Mika and Spo , al e na i e-
ly, as soon as he app oached an oes us emale. Hence,
we didn’ obse e any copula ion a emp by him.
Two o he p ime age males (K11 and K15) o med
small ma ing g oups o hei own du ing mos o he
peak u and si ed 7 and 6 cal es, espec i ely (Fig. 3).
Indeed, ep oduc i e success is pa ly ela ed o ma -
ing ac ic, which is highly dynamic, bo h empo ally
and spa ially, also among p ime males.
Mika - he “sma y”
Mika’s ac ic, s aying a ound he main g oup and
wai ing o he big chance, did pay o as seen by he
ab up shi in dominance be ween Spo and Mika
(Fig. 1), Ho being he ca alysa o . Indeed, o be
success ul males need expe ience; i.e. co ec assess-
men o own condi ion and i als s eng h combined
wi h op imal iming and alloca ion o hei limi ed
esou ces. The “ es os e one unaway” ac ic p ac ised
by Ho seemed no adap i e, bu he p obably lea ned
his lesson. Mika showed his expe ience also when in
cha ge o he g oup. He adjus ed his ac ic acco ding
o male-male compe i ion and emale a ailabili y by
combining g oup de ence wi h ending indi idual
emales. Using his i als as his ex ended “nose” and
allowing hem o ake pa in he “ o spiel”, Mika
p obably educed his se icing ime pe emale and
hence inc eased his access o ecep i e emales.
Body mass - no he only key o success
Males’ ank is clea ly ela ed o hei body mass in
polygynous ungula es (Clu on-B ock e al., 1988),
including eindee (Espma k, 1964; Røed e al.,
2002). As males’ dominance wi hin a ma ing g oup
gi es p io i y o access (Clu on-B ock e al., 1982),
he linea i be ween body mass and ep oduc i e
success (R2 = 0.56) is expec ed. The oldes male, Spo
was 6 yea s, and he s ong linea i be ween age and
success (R2 = 0.73) sugges s ha none o he males
had each senescence. Building up and main aining a
big and powe ul body is demanding. Indeed, males’
body mass e lec s physical ma u i y and hei capac-
i y o ake ac i e pa in he u . As soma ic e o
Fig. 3. Indi idual ep oduc i e success o all p ime (6,
5 & 4 y s old, black) and semi-p ime (3 y s old,
da k g ay) males and collec i ely success o he 2
(ligh g ay) and 1 (whi e) y s old males, espec-
i ely.

70 Rangi e , 32 (1), 2012
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inc eases wi h body mass (Fig. 4), he posi i e linea
ela ionship be ween e o (i.e. absolu e soma ic
loss) o ela i e e o (absolu e losses/body mass) and
ep oduc i e success ollows (R2 = 0.55 and R2 = 0.43,
espec i ely). Howe e , age speci ic analyses e ealed
no such ela ionship among young, no among semi
p ime males (bu only wo males we e a ailable).
Only among he p ime-aged males we ound a posi-
i e ela ionship, al hough a he weak (R2 = 0.21).
Bu , big guys and spende s can be lose s. Ho pu
much e o and igh ing spi i in o he u bu did
no ge any ou pu . Du ing his li e span (he was
slaugh e ed a e he u 2007) Ho ac ually ne e
si ed any o sp ing – could he be s e ile? Anyway, he
played a i al ole in he male dynamic. Excluding
Ho om he analyses he posi i e e o - success
ela ionship among p ime males imp o ed conside -
ably (R2 = 0.79).
Con ol is expensi e
The s ong linea nega i e ela ionship be ween ime
spen s anding ale and ea ing o Spo and Mika
du ing u (R2 = 0.68) sugges s ha hey aded
educed ene gy in ake wi h s aying ale and keeping
con ol, in line wi h he o aging cons ain hypo h-
esis (Pelle ie e al., 2009). In compa ison, he 1 y
old males spen 66% o aging. As a consequence hey
we e able o keep up hei body mass du ing u , only
losing 1 kg on a e age (Fig. 4). Indeed, hei sneaking
ac ic seems no o in ol e any big e o , whe eas he
semi p ime- and p ime-aged males’ ac ic, being sub-
dominan wi hin he main g oup ying o “chop o ”
single o small g oups o emales, was p e y demand-
ing (Fig. 4), as also con i med by hei inc ease ime
s anding ale and educed o aging ac i i y.
High s ake - big isk
The ul ima e goal o a p ime male is o con ol as
big a g oup o emales as possible du ing peak u .
Sup emacy, and hence p io i y o access o emales
wi hin a ma ing g oup is mani es ed h ough domi-
nance beha iou . Indeed, igh s a e he ul ima e
exp ession o assessing compe i i e abili ies among
p ime males. Al hough p ime males’ comba s pe
see seemed o con ibu e a mino pa o hei di ec
ep oduc i e spendings, igh s may lead o se e e
inju ies among polygynous ungula es (Geis , 1986).
Mikka’s le oo was ac ually angled in his own
an le s when igh ing o Ho (seen a he end o
ideo 1, jus a e Ho an away). P obably he go a
cu which led o a se ious in ec ion and his esigna-
ion in la e pa o peak u . Indeed, he winne may
pay his oll.
Conclusion
Reindee male ma ing s a egy is highly luid.
We ha e iden i ied ou main ( e e sible) ac ics;
1) Dominan ; con olling as big a g oup as pos-
sible and willing o ake he isks including igh s
o keep con ol as seen by Spo and Mika. While
con olling he g oup, ending indi iduals emales
wi hin he g oup coming in o oes us. 2) Wannabe;
includes p ime males willing o hang a ound as
subdominan s and wai o he chance o ake o e
ma ing g oups by igh s, as seen by Spo , Mika and
Ho . While wai ing o he chance hey p ac ise he
bu che ac ic (see nex poin ). 3) Bu che (Sa elli e);
ying o chop o and isola e small emale g oups
( om he main g oup) o single emales as seen by
he semi p ime males (K9, K5 & K3). 4) Sneake ; an
age speci ic ac ic, used by young male no able o
keep g oups o end indi idual emales a high male-
male compe i ion.
Aknowledgemen s
We wish o hank he Finnish Reindee He de s’
Associa ion and especially Mika Te onen and his c ew
a Ku uha ju Expe imen al Reindee S a ion o ca ying
ou he expe imen and he Finnish Game and Fishe ies
Resea ch Ins i u e, a he Reindee Resea ch S a ion in
Kaamanen o logis ic suppo .
Spo
Mika
K11
K15
Ho
K3
K9
Fig. 4. Rep oduc i e e o measu ed as body mass loss
among males du ing he u (Sep embe 6 h -
Oc obe 22nd) s ini ial body mass (Sep embe
6 h). P ime (6, 5 & 4 y s old; black big squa es.)
semi-p imed (3 y s old; da k g ay small squa es,
no body mass o K5 be o e u ), 2 y s (ligh g ay,
big iangles) and 1 y old males (whi e, small
iangles). Missing alues o one 3 y s and one
1 y old male.
71
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Video shoo ing
Videos a e shu by Léon L’I alien and Anne Lene Ho land.
Re e ences
B o-Jø gensen, J. 2011. In a- and In e sexual Con lic s
and Coope a ion in he E olu ion o Ma ing S a egies:
Lessons Lea n F om Ungula es. –E ol. Biol. 38: 28-41.
Clu on-B ock, T.H., Guinness, F.E., & Albon, S.D.
1982. Red dee . Beha iou and ecology o wo sexes. Uni e -
si y P ess, Edinbu gh.
Clu on-B ock, T.H., Albon, S.D., & Guinness, F.E.
1988. Rep oduc i e success in male and emale ed
dee . – In: Clu on-B ock, T.H. (ed.). Rep oduc i e success.
Uni e si y o Chicago P ess, Chicago.
Emlen, S.T. & O ing, L.W. 1977. Ecology, sexual Selec-
ion, and he E olu ion o Ma ing sys ems. – Science 197:
215-223.
Espma k, Y. 1964. S udies in dominance-subo dina ion
ela ionship in a g oup o semi-domes ica ed eindee
(Rangi e a andus L.). – Anim. Beha . 12: 420-426.
Geis , V. 1986. New e idence o high equency o an le
wounding in ce ids. – Can. J. Zool. 64: 380-384.
Holand, Ø., Røed, K.H., Weladji, R.B., Gjøs ein, H.,
Kumpula, J., Gailla d, J-M., Smi h, M.E., & Ni-
eminen M. 2006. Male age s uc u e in luences emales’
mass change du ing u in a polygynous ungula e – he
eindee (Rangi e a andus). – Beh. Ecol. and Sociobiol. :
59: 682-688.
Is a an, K. 2005. Va ia ion in male ma ing beha iou
wi hin ungula e popula ions: pa e ns and p ocesses.
–Cu en Science 89: 1192-1199.
Ma in, P. & Ba eson, P. 2007. Measu ing Beha iou :
An In oduc o y Guide. Camb idge Uni e si y P ess,
Camb idge.
Mys e ud, A., Bonen an , C., Loe, L.E., Lang a n, R.,
Yoccoz, N.G., & S ense h, N.C. 2008. The iming o
male ep oduc i e e o ela i e o emale o ula ion in a
cap ial b eede . – J. An. Ecol. 77: 469-477.
Pelle ie , F., Mainguy, J., & Cô é, S.D. 2009. Ru -In-
duced Hypophagia in Male Bigho n Sheep and Moun-
ain Goa s: Fo aging Unde Time Budge Cons ain s.
– E hol. 115: 141-151.
Pembe on, J.M., Albon, S.D., Guinness, F.E., Clu -
on-B ock, T.H., & Do e , G.A. 1992. Beha io al
es ima es o male ma ing success es ed by DNA in-
ge p in ing in a polygynous mammal. – Beha . Ecol. 3:
66-75.
P es on, B.T., S e enson, I.R., & Wilson, K. 2003. Soay
ams a ge ep oduc i e ac i i y owa ds p omiscuous
emales’ op imal insemina ion pe iod. – P oc. Royal Soc.
London, Se ies B 270: 2073-2078.
Rops ad, E. 2000. Rep oduc ion in emale eindee . – An.
Rep od. Science 60-61: 561-570.
Røed, K.H., Holand, Ø., Smi h, M.E., Gjøs ein, H.,
Kumpula, J., & Nieminen, M. 2002. Rep oduc i e
success in eindee males in a he d wi h a ying sex a-
io. – Mol. Ecol. 11: 1239-1243.
Wade, M.J. & Shus e , S.M. 2004. Sexual selec ion: ha-
em size and he a iance in male ep oduc i e success.
– Am. Na . 164: 83-89.
Manusc ip accep ed 16 Ma ch, 2012
72 Rangi e , 32 (1), 2012
This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License
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Appendix 1
Me hodology
The s udy was ca ied ou a Ku huha ju Reindee
Resea ch S a ion (69ºN, 27ºE) in no he n Finland,
2007. Be o e u he he d was ounded up and all
animals we e weigh ed (Sep embe 6 h). All males
(excep se en 1 y old) we e i ed wi h adio col-
la s, whe eas he es o he he d we e equipped wi h
numbe ed colla s making indi idual iden i ica ion
possible. Blood was d awn om all animals. The
expe imen al he d was eleased on o he u ing
g ound Sep embe 8 h.
We acked as many as possible ma ing g oups (i.e.
agg ega es consis ing o a leas one emale wi h a
leas one male p esen ) daily om Sep embe 16 h o
Oc obe 15 h, gi ing p io i y o he p ime and semi-
p ime males. All animals associa ed wi h each g oup
we e eco ded and he dominan male assessed based
on obse ed u ing beha iou and agonis ic in e ac-
ions be ween males. We eco ded males’ ac i i y
(i.e. es ing, walking, s anding ale and ea ing) and
di ec u ing beha iou (i.e. lehmen, ipping back-
legs while u ina ing, ubbing an le s, in es iga ing
emales, ending, cou shipping, chasing and igh ing
i als and he ding emales) e e y 15 seconds o 15
minu es based on he ocal animal echnique (Ma -
in & Ba eson, 2007). All ocal obse a ion pe iods
con aining he ac i i y class es we e omi ed. In
addi ion, males ocaliza ion (numbe o g un s) o e
a 5 minu es pe iod was eco ded a he end o he
ocal obse a ion pe iod, as a measu e o exci emen
le el and hence he in ensi y o he u . The dominan
male wi hin he g oup was he i s o be chosen o
ocal obse a ion. The ea e subo dina e male(s) p e-
sen was (we e) chosen. The dominan male wi hin
a g oup could ac ually be obse ed se e al imes
du ing a day. We he e epo a e age alues o daily
obse a ions.
All copula ion a emp s, de ined as moun ing o a
leas 2 seconds we e eco ded. Consecu i e copula ion
a emp s o he same emale by he same male wi hin
he same day we e coun ed as one a emp . Fu he
he u was di ided in o p e, peak and pos u . Peak
u was de ined as he sho es pe iod whe e a leas
90 % o all copula ion a emp s we e obse ed. Ac u-
ally 22 ou o o ally 24 copula ion a emp s in he
main ma ing g oup we e obse ed du ing peak u
(Oc obe 2nd – 9 h) (Fig. 2). The p e u and pos u
we e de ined as one week p io o and he six days
a e peak u , espec i ely.
The he d was again ounded up Oc obe 20 h and
all animals weigh ed (Oc obe 22nd) be o e eleased
o hei win e ange. Ele en elde ly emales we e
slaugh e ed in Oc obe and Decembe 2007 and wo
died du ing he ollowing win e . P io o cal ing in
2008 he emales we e ounded up and con ined o a
smalle cal ing a ea. Jus a e cal ing he cal es we e
weigh ed and blood samples d awn o pa e ni y anal-
yses using mic osa elli e ma ke s (Røed e al., 2002).
Fou emales did no gi e bi h. Al oge he 70 cal es
we e sampled and 69 assigned o one o he males.