Shit happens - a glimpse into males’ mating tactics in a polygynous ungulate - the reindeer

65 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 Edi o in Chie : Rol Egil Hauge ud, G aphic Design: Lundblad Media AS, www. angi e .no 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 This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License Edi o in Chie : Rol Egil Hauge ud, G aphic Design: Lundblad Media AS, www. angi e .no 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 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 Edi o in Chie : Rol Egil Hauge ud, G aphic Design: Lundblad Media AS, www. angi e .no 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 68 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 Edi o in Chie : Rol Egil Hauge ud, G aphic Design: Lundblad Media AS, www. angi e .no 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 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 Edi o in Chie : Rol Egil Hauge ud, G aphic Design: Lundblad Media AS, www. angi e .no 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 This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License Edi o in Chie : Rol Egil Hauge ud, G aphic Design: Lundblad Media AS, www. angi e .no 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 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 Edi o in Chie : Rol Egil Hauge ud, G aphic Design: Lundblad Media AS, www. angi e .no Video shoo ing Videos a e shu by Léon L’I alien and Anne Lene Ho land. 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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.