Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. estl onesrtge hf h aac fitreulcnitfo ae ofemales Xiang to Zuofu males from conflict intersexual title Short of balance males the from shift conflict intersexual counterstrategies of Versatile between balance interactions the dynamic during shift how as can show males societies findings multilevel to Our in beneficial conditions remain. likely the who is how females. mothers which and another to the infants, be to attacking of can compare transfer cease offspring females replacements mothers’ and future males for male males sire some opportunity during can transfer, an present they mothers’ provide infants tenure to and of longer only infanticide response mortality to delay In in crucially lead difference unit. defenses which no joint social infants, Female and the infanticide with periods. to other transfer succumbing to and infants mount defense to 31 joint able whether of female are unclear females 4 mating, remained that extra-unit has reveal roxellana including it Rhinopithecus infanticide, counterstrategies, in male observations versatile long-term against Detailed counterstrategies develop effective. are to counterstrategies predicted are females primate Although Abstract 2020 4, August 3 2 1 Xiang Zuofu females conflict to intersexual males of from balance the shift counterstrategies Versatile eta ot nvriyo oety&Technology, & Forestry of University South Biology, Central Conservation and Ecology Evolutionary China of P.R. Institute 410018, Hunan, Changsha, Nanlu, Shaoshan 498 Xiang Zuofu author: *Correspondence China 4 China 100101, Beijing, 3 2 China 410004, Hunan, 1 affiliation: Author nttt fZooy hns cdm fSciences of Academy Chinese Zoology, of Province Institute Hubei Park, Technology National & Forestry Shennongjia of University South Central e a fAia clg n osrainBooy nttt fZooy hns cdm fSciences, of Academy Chinese Zoology, of Institute Biology, Conservation and Ecology Animal of Lab Key etrfrEclec nAia vlto n eeis hns cdm fSine,Knig 650223, Kunming, Sciences, of Academy Chinese Genetics, and Evolution Animal in Excellence for Center hnogi ainlPr,HbiPoic,SennjaFrs itit ue 441 China 442411, Hubei District, Forest Shennongjia Province, Hubei Park, National Shennongjia olg fLf cec n ehooy eta ot nvriyo oety&Tcnlg,Changsha, Technology, & Forestry of University South Central Technology, and Science Life of College oee natcd nsu-oe monkeys snub-nosed in infanticide Lowered : 1,* 1 agYu Yang , agYU Yang , 1 u Yao Hui , 1 u Yao Hui , 2 igagHu Qinglang , 1 igagHu Qinglang , 1 igLi Ming , 1 2 n igLi Ming and , , * 3,4 3 Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. ta.21) u nisl,fml oltoaydfnehdbe osdrda nffciebcuei a only can it because ineffective as considered been the had on Zipple defense 2000; improved depends coalitionary Schaik having van likely female from & conflict itself, (Nunn benefit in sexual infanticidal might but be the females might 2017), in their species, that al. males balance investigate gregarious et against in This studies defense example, coalitionary few For react. and species, detection might system. new various males mating a in and how joining reported social and or been if male, have ousted and the counterstrategies effectiveness male, with female protector 2012). transferring a though al. male, or Even et infanticidal females Roberts the other 2012; and male, with have (Palombit infanticidal unit coalition unit they the in social during social from changes, or the offspring spatially own dependent leaving rank it have their by male separating females on or by If or males offspring take-overs infanticidal 2012). their counter-attacking as pregnant al. defend by are et such sometimes a females (Roberts they occur, generating If that pregnancies situations, to certain their 2014). 2019), these the Huchard terminate be likely reduce & to cannot is al. Lukas shown might found dominants 2004; infanticide Females et been been future MacDonald, have when that Ma & infanticide. (Wolff such that situations 2013; male own males, behavior during their from many of not al. with loss is range mating et infant offspring a by given avoid (Yin infanticide exhibit to males of primates males counterstrategies risk by the female overall be among killed turn, change to In be status suggested by resident might or females. or success infants rank infants reproductive to frequent unweaned his unrelated costs with of increase killing considerable species to 50% some 2016): killer In to the al. sooner. up permitting et infant Yao mother, next adult 2012; the her by Palombit of siring young interval unweaned 1997; of inter-birth (Hrdy infanticide the example with shortens primates, ubiquitous non-human most in conflict the sexual males switch of dynamic forms most potential many However, and are counter-strategies. interactions There 2015). and of Palombit strategies understanding successful 1998; al. our of either (Ebensperger limiting to use et males sex, leading the (Chapman single of conditions in a adaptive the expense on determine the are focused to at have females females gaining studies males of to females of perspective harm to the perspective involving or from the resistance case strategies from some which expense studied in the been under or at frequently 2003) fitness creating conditions & has their the (Chapman diverge, conflict maximize coevolution to unravel Intersexual often selected antagonistic to 1972). are males in (Trivers sex mates result each and of their can females individuals of conflict where how of Such 1996), show Rice strategies 2006). 1996; (Parker, findings can Partridge reproductive sexes Our societies the the multilevel between species, in remain. conflict females. conditions reproducing to who as the males sexually mothers how males from In and to the conflict intersexual be beneficial of of can likely offspring balance females is the future and which shift unit. males sire infants, social between attacking no can another interactions cease and joint they to dynamic Female infanticide males transfer tenure to some mothers’ periods. longer succumbing transfer, for other infants during mothers’ opportunity to 31 to an compare of provide response replacements 4 and male In joint infanticide only during female delay to present mating, crucially infants lead defenses extra-unit of which including mortality infants, counterstrategies, in the versatile difference with mount transfer to in and able observations defense are long-term remained females Detailed has that it effective. infanticide, reveal are male against counterstrategies counterstrategies whether develop to unclear predicted are females primate Although Abstract analysis. paternity the counterstrategies, conducted female ML personality infanticide, and polymorphic male ZX society, conflict, and observation, Sexual field : the Keywords performed QH and ZZ YY, HY, ZX, Authorship: ( Li M ([email protected]) Xiang Z 86-731-5623498 Fax: 86-731-5623392; Tel: [email protected] XadM ocie h eerhdsg,aaye h aa n rt h ae jointly. paper the wrote and data, the analyzed design, research the conceived ML and ZX ) 2 hnpteu roxellana Rhinopithecus hnpteu roxellana Rhinopithecus multi-level , Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. takatrlcaigfml rnfrigwt nendifn f()ml eue r ogrta females’ tenures than longer have longer might infants. are attack unweaned tenures without to attack infant refraining male consistently unweaned the or (i) who with tolerating sire tolerate if males from to who infant than benefit male likely unweaned would (ii) more with males and/or are intervals, that transferring success birth predict males reproductive female we resident immediate lactating OMUs, benefits. as we need after their Furthermore, they their infant, attack in because females success. unweaned and aggressive of reproductive the counterstrategies offspring be future towards next to females’ for aggression, likely wait the more or cannot are to and tolerance OMU. joint males response larger behaviors, female older in different that in because efficient two predict attacked males more have or they be of males killed also where strategies Considered might being or investigate vigilance of leader and size risk next detection larger male less We male that early the lead predict and/or ousted we with would stronger the defense, infant) mated be joint with the might previously through either defense from except offspring have protection, defend attack (members they able male unsuccessful OMU are where an seek of females OMU suffer (iv) to relatives; new infant OMUs with female unweaned a have original or the into their OMU) if unweaned immigrating leave their and (iii) would or mother over OMUs); the females take these for would male, protection into offer who new immigrate could unit, with they who including OMUs, all-male if OMU, or other the infant their of with of in males replacement leader (resident males the male protectors than adult potential following other pregnancies (unassociated males with their killers copulate terminate particular, potential females In would (ii) times. females aggression; other male pregnant during and without present (i) take-overs those that male of and difference during replacement no expect population to show male behavior we would of free-ranging counterstrategies infants times male unweaned a during female of present mortality and in infants and of between infants effective female effectiveness and or versatile of the be fetus observations would counter-strategies of for detailed fate support 15-year, the the a of investigate on female firstly and based which infanticide we partners infanticide male study, social offer of this expression might close the monkeys among In on snub-nosed remain interactions in but facilitating social flexibility 2019), of dispersal social role and of al. the the counterstrategies. among costs investigate et Therefore, association low to Zhang spatial opportunity have attacks. the 2009; an potentially against addition, been females support al. In have provide that et 2016). the paternities might means (Qi have al. extra-unit also units et they and Yao between units as 2010; OMU, transfer across paternity own al. kin obscure et their of to (Guo outside distribution chance species males with the this with transfer in have contact confirmed contacts mothers females social regular defense, that of interactions nestedness joint for means The detailed female opportunity monkeys but unclear. counter- snub-nosed remain i.e. 2020), counterstrategies, one golden these females, as al. to in and response et proposed males’ males (Qi been and between infants monkeys has their switch survive snub-nosed promiscuity replacement dynamic golden position female male potential dominant Recently, of in the times and infanticide over male. the male take during new males present against the infants when strategy unweaned by observed most attacked been but has being (OMU) 2016), (Xiang Infanticide without unit band al. 2020). social male et larger (Yao one al. one OMUs of as et in together primarily Grueter forage consisting 2014; and units associate al. OMUs social et such multiple core and in females, reside multiple with females which in behavior sex. society, their opposite multi-level ( of the monkeys aspects of females. snub-nosed different individuals flexible these Golden change in the of to dynamic of example, particular opportunities reproductions behavior For reproductive be the the future future to have on during infants. females gain might depending control and attack might individuals males mating some male to between where and this stopping conflict societies, tenure infants, sexual from attacking his expect benefits would stopped retain we gain who can Accordingly, might male he males a transfer if leave to turn, opportunities expected not order in be in would might females if, counterstrategies females Female other effective if by species. more tolerated territorial other be in be from limited or protection might to societies; paternity; which uni-male confuse females group/unit, to in new which order hnpteu roxellana Rhinopithecus 3 r sa ooie nei oCialvn na in living China to endemic colobines Asian are ) .roxellana R. ehpteieta females’ that hypothesize We . Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. eiigi h l-aeui,ad(i rqece fetaui ouaino hs eaeivle nmale in involved female those of males copulation or extra-unit units of one-male frequencies extra-unit any in (ii) (i) residing and whether identify males unit, to all-male established used was the and test in females chi-square residing between A occurred unit. social primarily were different copulations dyads a study in ”sister-sister” in resided half farther dyads five the female and of 2006, analysis social dyads relatedness since Data ”sister-sister” the data. The OMUs full genetic within into six the residing loci. born and on male dyads based infants more the ”mother-daughter” identified observing or male all by two extra-unit categories: determined at an three were into to excluded divided assigned was was levels was conception population confidence paternity our the with which of 2007) from during in reliability band al. unit cases the The et into all (Kalinowsk born in (2014)]. CERVUS3.0 offspring software and 99 al. the the et with for Yang Paternity determined of was and 99.1%. samples 2018 was blood (2014) matched agreement to end with the 2007 al. patterns its allele individuals: et comparing on captive by Xiang five glue confirmed see of was with samples D10S611) protocols hair exclusion D6S474, pole from Paternity DNA D6S493, D5S1457, results a genotyping the 2015). D14S306, D2S442, using on D6S1056, D1S533, al. D7S2204, D10S1432, et details by D7S1826, (Guo (D13S321, [for Hair obtained D1S1665, individuals loci were D1S1656, target 2015). samples microsatellite D10S676, the – different while of D7S817, al. gloves) approach sixteen back et (with using to the (Guo hand established with difficult was by contact band were in directly focal brought who them the was plucking – which in by infants males obtained young adult period, were study of mature bouts individuals the during aggressive reproductively adult born otherwise all of infants all attack, samples from from collected of and were set mothers, roots have intact first their with to hairs the attempts. were defined analysis, during infanticide paternity females was the unsuccessful male For the attempt as the and infanticide scored by male successful were male new killed infant A previous the was the the between infant towards male. et when copulations the previous occurred (Xiang when when have the or began to occurred et AMU by event decided (Yao the disturbance an was takeover joined once without male the and previous sampling observed of unit the year all-occurrence original of the continuous the replacement of via any left final November) recorded as The (late extended were well 2019). season sampling events as animal conception) al. All infants, focal (potential with This 2016). mating mothers sampling. the al. and animal of infants focal end a females, the in pregnant to observed abortions, on were births, events attacks of infanticide male instances abortions, using place, All observed taken 2020. were had March event takeovers and male 2006 January and between copulations summer-autumn in 0700-1900 three or comprising to individuals 2006 ˜30 January collection from in Data in increased lip individuals) has upper band adult the ( focal of (five physical AMU the sides nipples, one AMU of of female’s both and associated size set a on 50 of The an nodules unique and size fleshy and 2013). a and 5 of OMUs shape al. present on between the et are reserve based distances scars, (Yu which or identifiable and males at flanges injuries adult granulomatous individually researchers basis of of were evidence of daily shapes color, members the a presence pelage and adult size, on the body All made ignore as monkeys’ 2019). were habituated such to the features, al. observations became started affected et Therefore, soon monkeys likely (Xiang time. they Provisioning the However, m the km. fleeing, process. of 1 the of most ˜ in instead staff carrots, of early apples, and radius especially seeds, a humans, activities, provisio- pine with to not daily lichen, Zhang area When and included 2013; an sites/platforms. behavior and al. feeding across day social different et ranged per (Yu at monkeys provisioned times ecotourism the were three of monkeys ned, to purpose the provisioned. two the peaches; partly provided and for but was oranges, 2006 free-ranging, Food in is 2019). monkeys al. population the et study provisioning The began China. staff central Reserve Park, National Shennongjia subjects in and sites Study METHODS AND MATERIALS > 0 rlxdcndne and confidence) (relaxed 80% > xr-ntcpltosadetaui aent eejde ohv curdi h aeor mate the if occurred have to judged were paternity extra-unit and copulations Extra-unit ecletddt nadiybssa itne 0mfo 80t 80i winter-spring in 1800 to 0800 from m 50 - 5 distances at basis daily a on data collected We ee dl ae)i eebr2019. December in males) adult seven ecnutdtesuya aoga (31 Dalongtan at study the conducted We > 5 src ofiec) hs neecsmthdecuinstandards, exclusion matched inferences These confidence). (strict 95% 4 dlibitum ad apig oee,oc aetakeover male a once However, sampling. ° 9N 110 29’N, > 0idvdasi v OMUs five in individuals 90 ° 8E lvto:20 m) 2200 elevation: 18’E; Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. aigwt h eietml ftedsiainOUit hc hytaserd u oextra-unit no but transferred, they observed which been into previously OMU had OMU destination #16) another cases. case the two to in these of transferred Yb for and male #6, identified OMU were resident Case original paternities in the their (Ln left with mothers OMUs who mating two the infants in replacement, unweaned offspring male with any following mothers of fathers nine males the the new not Of All were accordingly (6.25%). and births AMUs, 99 (309 from in entered. units identified came they were one-male OMUs OMUs, over other ( other took events) in in who (8 males males unit resident exclusively all-male involved the almost which paternity, in involved males these than and rather events) (6.7%), matings extra-unit were 0.7102, = mating Z-Score Extra-unit 19/92, infanticide vs male (8/33 no against periods is counterstrategies stable There Female present those those times. and and other the replacement replacement at male and 0.3408, 0.05). male present of = infants of times Z-Score those the during times 17/90, and between present vs during infants replacement rates (6/31 the periods death male between stable of of rates present difference death times of the during difference because compared present (potentially significant We winter infants in reasons + disease). unknown and fetus for dying temperature replacement low Therefore, male of of events. times takeover during male present were of who ( outside replacement male pregnancies following 117 abortions p of of stable increase total significant to a a compared is during there replacement observed male were abortions during Two infants and unit fetus social ( of another male the attack, mortality most periods to new throughout in of transferred infanticides, the survived by male, difference set four tolerated ousted have The first of were the to the except or with known attack, by stayed Therefore, unsuccessful are mothers killed an their replacements (#8). were (15/29) after because attacks infants #11) male either unweaned and repeated during season remaining #8 after present mating all mothers killed (#2, (27/31) of their was infants infants half infant and three unweaned Although more only them survived. one of males, and and Two new male new by ousted the replacements. attacked the by male male were with attacked ousted following unit not with present the were stay left were females to pregnant infants returned other unweaned one one offspring. All and Thirty-one #3), their (#11), (#17). and to male (#1 offspring birth new gave abortions her events, the and two weaned by replacement males in and killed male resulted birth was subsequently during gave which newborn present and males, the were new but females by birth pregnant attacked gave and were Eight 2006 which January of #18). between four and males adult (#7 15 #18, offspring and - unweaned OMUs #1 16 (Case involving 2020 events March replacement fortunes male their 18 and observed We infants unweaned females, pregnant male tolerance replacement, the Male between tenures attacks, the suffer in not (iii) between did RESULTS and infants infant) which tenure, the in males’ except infant those and male. versus interval (members or aggressive males birth OMU and fetal by females’ killed of if tend between or size (i) (ii) females attacked if (i) were identify pregnant independent infants differences two-sample to if which significant A (ii) used in were OMUs was circumstances, replacement. there other test male whether than score following establish higher pregnancies Z their are A terminate replacement involved. to male not following those rates than death higher are replacement < .0) wnytreuwae natdah eeosre uigattlo 2 its ihtoinfants two with births, 121 of total a during observed were deaths infant unweaned Twenty-three 0.001). nttl ,9 ouaineet eeosre.Ol 1 fteecplto events copulation these of 317 Only observed. were events copulation 4,697 total, In al S1 Table .Toeet curdwe ofmlswr rgato aigfor caring or pregnant were females no when occurred events Two ). 5 p χ 2 > 182.00, = .5,adbtentefts+ifnspresent infants + fetus the between and 0.05), i 1 Fig i 2 Fig p and < .0) ee ae fextra-unit of cases Seven 0.001). / s217 -cr 3.6212, = Z-Score 2/117, vs 2/8 , al S1) Table . t ts a sdto used was -test p > Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. ymls(/1.Mlsi ungi rmcagn hi w taey stlrtn nendifnsmeans infants unweaned tolerating as killed strategy, were when own take-overs infants their to during present changing further OMUs infants from any of gain other in attacking number turn to small stop in as a transfer Males and/or only effective, (4/31). thus successfully infant males appear and other dependent to by infants, counter-strategies with with the able These left defense tolerate are mothers joint males. would some and form infanticidal males attacks, males, from most male extra-unit offspring response with against unweaned living mate OMU that their can same suggest remove females years the that fifteen in means across females male-takeovers society during multi-level responses a female in of observations detailed The ( offspring current longer her DISCUSSION months, wean 39.4 successfully average with to on female benefits ( lasted a reproductive until males take-over takes future longer after it increased wait tenures months 62, gain to male 23.5 = to needed female average) (i) likely they (on lactating emigrate: are 14-16), the not when but # than did or females, and who #13) these #9 females and of (#5, those offspring #12 attacks next (#10, further the attack from siring tolerant any refrain than males without older the of infant not transferred unweaned are set by favoured males first tolerated aggressive be the they to benefits, (Case if expected means fitness be immediate emigrated sooner might on females infanticide mother ( more Although lactating victims’ males rely #17). as might and of #6 who benefits offspring (#4, males, lost offspring older next the completely the kill they not repro- sired However, did increased attack they #11). obtained because and they bring aggression, infants #8 would of which #2, killing strategy tolerance, from a or success adopted aggression males ductive either When strategies, success. two showed reproductive males different another new benefit replacement, joined its male collectively After and who infant females two unweaned towards of mating sets strategies observed two Males’ been another (iii) had relative and or who clique. female unit, females offspring close a destination two one as their of (ii) least OMU male at joined, on had resident they attacks who unit the returned females unsuccessful new with or five the the (i) OMUs in included to other sister-sister) OMU (mother-daughter, response existing to already transferred in an (11/31) into male ( immigrated mothers (ousted) OMU of the old third in the offspring a with Nearly stay infants to with transfer emales’ F which ( ( in infants observed . OMUs killed were in never attacks smaller males no was which which size in OMU in those as OMUs infants), in except depend of than to individuals p most observed appears of OMU, defense been number joint another the (the had female in OMU to attacks of since of effectiveness transferred and size The subsequently unsuccessful, weaning. the attacked to being on survived were attacks 94.1%) infants male (16/17, 3.38 initial whose infants the these was mothers of cases defense (17/20) of collective 85.0% in majority in usually resulted he involved females male, females of the time. adult behavior at collectively some shouting of female for and number non-lactating females facing cases, the The females’ from few to away response a stayed In the In (and the and away. at In unit mother). quickly it shouting keep the started retreated same male. and already (including the the female perpetrator others at of other the by loudly while chased females joined infant vocalizing other her being after started seizing she, Then, before subsequently still as male and was coalitions mother ( perpetrator. mother of the the male the instances (10/15) to new at remaining 66.7% ran shouting the of juveniles) initiator started against also the infant, females sometimes was her other attack seizing under with was joint firstly infant to whose observed female were lactating 59/71) females (83.1%, other OMU with defense Joint < .0) n h ieo Msi hc nat idfo natcd a losgicnl mle than smaller significantly also was infanticide from died infants which in OMUs of size the and 0.001), p t t 2.31, = < 1.93, = 0.001, df df i 4a Fig 9, = 14, = i 1 Fig p ,ad i)tnrso oeatmlsaesgicn ogrta hs faggressive of those than longer significant are males tolerant of tenures (ii) and, ), p < .78.O h te ie hnmlsaotdasrtg ftlrne which tolerance, of strategy a adopted males when side, other the On 0.0738). = 0.05, .Tecnet ftenn ohr ihdpnetosrn h successfully who offspring dependent with mothers nine the of contexts The ). ncsso aeatcso nendifns otfmlso h same the of females most infants, unweaned on attacks male of cases In i 4b Fig i 3b Fig ). , t 2.53, = 6 df 29, = p ± < .9(mean 0.89 0.05). i 3a Fig ± , D.Tedefensive The SD). t al S1 Table 4.57, = t 3.55, = df .The ). 26, = df Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. ae(ry19) salsiganwui ihteose ae swsosre nfu ae ee is here, cases four in observed was as male, killer ousted potential the a from with offspring unit unweaned new their remove a can Establishing mothers be 1997). unit also (Hrdy might social male detection their adult risk from more and transferring because vigilance By potentially coalitions, units, opportunities female-female larger stronger the 2012). in to have (Palombit pronounced lead efficient might more Waal can more was they OMU (de effect until exhibit an offspring This to alive in females unit. killing females offspring for their delay from their from crucial of keeping males the transfer create to 85% particular prevent might nearly it in completely monkeys in in counterstrategies, snub-nosed to considered offspring in own, other been that unable their their suggest sometimes killing be findings on has from our attacks might attack 1997), male males male females against preventing a defenses in because joint resist effective ineffective this female to were although In unable Therefore, that 2012). instance. defenses Palombit be that joint 1998; might formed suggested (Ebensperger females been females physically while has attacks infanticide that, it prevent primates, find polygynandrous to in we some unable males study, in and be females does would between it mothers dimorphism mate unit as that pronounced to frequently further this is even females from there potentially for As infanticide came opportunities of OMUs limiting risk over presumably the 2014), taking reduce societies. al. males to mated males outside rarely et and all these very them (Xiang towards females though aggression with identified; monkeys contrast, intense even were snub-nosed In show collectively 2020). resident golden unit, paternities OMUs al. a in no all-male multiple et from with (Qi there the males unit study resident although in recent a because mating a residing into Yang in observed accepted males 2010; reported been were al. also with et previously was females (Wolff Guo case where had 2007; father one cases she Zhao another their two & infanticide, whom be extra-OMU (Li observed prevent with OMU and might we to other male mating Furthermore, they in strategies extra-OMU 2014). if residing monkeys, by effective al. offspring males snub-nosed mating most and et killing In females Multi-male the 2014). from between Huchard counterstrategies. of refrain occur & multiple paternities males one Lukas on that 2004; be such rely MacDonald, to & paternity and indicated obscures partners flexibility been it the social have has as they of mammals if infanticide range in male a against females offspring to with of their risk interact not defend to the to is able females reduce best strategy to because appear counter-strategies primates evolutionary low Female other relatively and best many is August have the infanticide females late Therefore, male infanticide. pregnant between male of counter-strategies. the breeders occurring not risk because effective seasonal conceptions could gestation the and most strictly females terminate Second, First, versatile and are The 2017). reasons. May more al. monkeys August. several to have et snub-nosed before to March (Xiang golden pregnancy from due November terminate evolve because occurring early to birth to abortions most failed females after with likely for shortly has benefit conceptive offspring, effect additional be their Bruce no and the females is 2017). rather towards monkeys, there al. aggressive feticide snub-nosed et are selected (Zipple golden males sexually baboon In new savannah takeovers. of male the in result to shown where the response been instances obligate be has in of an as might effect studies not reported monkey Bruce is rates other snub-nosed the population death In study golden fetal than than in replaced. in lower loss much is abortions fetal counter-strategy is that male Therefore, (25%) This suggesting death dominant ( studies, fetal 2012). a those of gelada al. rate in after the the et However, weeks primate, takeover. effect, (Roberts Hackl male the wild following Bruce 1983; birth in a (Berger the highly its including terminated effect exhibit is mammals pregnancies after Bruce it to wild of killed as argued other strategy 80% be adaptive in been with will an have confirmed as offspring females been males female’s has unfamiliar males, a to by exposure that infanticide after likely of pregnancies shifting terminate threat in in they them males. important the where towards with be to or offspring might females response siring constraints towards of social In either chance that conflict more suggest sexual a findings of have These balance would tenure. the males longer tolerant with and future him, the with stay mothers their ne rod19) oeta 0 rgatfmlsidc abortions induce females pregnant 90% than more 1999), Arnold & ¨ ander 7 hrpteu gelada Theropithecus ) Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. pce She l 04.Atlrn aewudovosyoti uuerpoutv ucs hog longer a through confronted success reproductive when many future population in obtain predator the obviously a to would resources to male benefit patch fight tolerant interspecific with also A in ( competed but 2004). tenures bolder when sleeping) al. mean OMU et for also (Sih his conspecific better species to in or aggression females fruit more the more because to predator with benefit tree infanticide, only following by one infants not success (i.e., unweaned reproductive may in towards immediate male personality personalities obtain resident male larger this obviously of of would one and kinds male personality as two aggressive animal polymorphic observed An together allow have replacement. reproduction, forage would we male study, complexity and future this social snub-nosed coordinate In and In 2020), band. OMUs 2007). al. current the multiple time al. et where between et (Grueter across (Wolf society, band wait trade-off behavior population multilevel social polymorphic to a in the in of span differences explain that monkeys adaptive individual life insight golden an explained consistent given enough the dependent be be as might not could the on it termed personalities have kill that Based often argued might to We contexts. which likely males. they personality and younger not fitness, those male’s are as their the female males by lactating increase older of obviously why cycle reproductive for would another explanation it tentative although a 4/31). infant efficacy provide the (12.9%, want on monkeys influence we an snub-nosed Finally, have golden would replacement infanticide. system than male against mating higher following or counterstrategies hanuman infant organization female also of social of of which society that rates implicate (8/37) multi-male death might and 21.6% 1997), data uni-male These and Borries society Asian In 1987; (11/29), uni-male Sommer 2013). some 37.9% an 1984; al. were in al. in et observed et example, (Yin mortality (Winkler takeovers For langurs the male societies. ma- after than multi-male involved infanticide lower or infants possible ( much single-male those langurs in also for mortality white-headed live was infanticide base-line of females from population the which mortality study to in Furthermore, replacement the colobines 2). no male in (Fig to during replacement leading groups coun- present le transfer, effective stable infant and and in of defense versatile infants mortality joint mount of the mating, to his female. in extra-unit able that increase difference including are of significant can infanticide cycle monkeys he male reproductive snub-nosed against because another females an as terstrategies least also accept that females at but only confirm for of not we her, dominant offspring might Additionally, with remains OMU future he mated an sire if previously of current to success he male their reproductive likely resident kill because future are a not female Similarly, males even do paternity. lactating males and they extra-unit tolerant immigrating when males, of cycle, refrain band aggressive probability reproductive they lose study low female those if the the the would fitness than in than higher tenure they longer case gain longer significant to the that are have likely in tenures means are is male its males females It average because the if infant. on offspring adult From success unweaned that of too. the reproductive appears attacking it behavior, current emigration OMU, from increase male unit, the might influence in social infanticide remains to mother a While appear completely. over transfers to opportunities taking female proximity reproductive males with close of associated in costs perspective be (Grueter low to addition, areas relative same In units The the form infanticide. facilitating within leaves of resting 2006), and and risk lichens (Li 2020). feeding, ubiquitous high al. travelling, year because is et through the this low attacks there is band, predatory throughout when species reduce the constituent potentially this refuge int that dietary in male a means be member major resident infant and This band other unweaned a join the 2015). with her to among al. experienced them and competition et sexual help her food have (Guo might of offer females which band dispersal also OMUs, Some secondary natal might neighbor units. compared and their in these reduced natal within residing in be both relatives units tolerated to and female social likely golden have sub-adults, other in are frequently as as into unit females a disperse such new occurs defend already where societies a multi-level mostly females that unit into In females Some groups 1997). social transferring societies. with al. of established et other saturated costs already (Steenbeek to not potential offspring an is the the into habitat monkeys, kill transfer to i 4b) Fig n oooieptriyb trcigfmls eiec.Teeoe grsiemlsare males aggressive Therefore, residency. females’ attracting by paternity monopolize and rcyihcsleucocephalus Trachypithecus 8 ,aleee nat one hn6mnh idof died months 6 than younger infants eleven all ), Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. u,S,J,W,L,M,Cag .&L,B 21) h aigsse fteScunsu-oe monkey snub-nosed Sichuan the of multilevel system primate mating a The of (2010). formation B. the Li, in & H. kinship Chang, of M., role Li, The ( W., Ji, (2015). S., B. Guo, Li, & W. Ji, K., society. Z., Huang, Xiang, S., M., Guo, Li, T., Bergman, D., sociality. Zinner, mammals. animal X., in Qi, infanticide C.C., Grueter, to counterstrategies and Strategies conflict. 321–346. Sexual (1998). (2003). L.A. L. Rowe, evolution. Ebensperger, & for J. ( fuel Bangham, as G., langurs conflict Arnqvist, Hanuman Sexual T., Chapman, of (1996). L. groups Partridge multimale & T. breeding Chapman, seasonally in Infanticide horses. wild entellus in (1997). factors C. social and Borries, abortion Induced material. supplementary (1983). electronic J. the Berger, of part as uploaded been REFERENCES have article this supporting datasets The availability interests. materials competing and no Data have they that declare authors The State the and China (2016YFC0503200) interests Pro- of China Competing Research Foundation of Priority Science constructive Strategic Program Natural China. his of the National R&D for Administration by the Lukas Key Forestry Supported (XDB31000000), Dieter 31821001),National by Sciences Dr 31670397, of supported and (31870509, Academy was help Clutton-Brock Chinese and study Tim Lu of This Professor Lang gram Wei, paper. thank Li this We Zhang, on field. Bo comments Liu, the Ruoshuang in Chen, data Hanlong Chen, collect Haochun Fan, Penglai thank We (IACUC) Committee Cen- Use by and approved Care Acknowledgments and Animal evaluated Institutional were Technology’s and & requirements #2012-018). Forestry legal (protocol of obtained Chinese University the male South to the tral adhered case, methods case. first research later the All in were In females infants lactating #17). two all (Case result lost which emigration he Ethics infant however, females dependent success, lactating the reproductive instance, two attacking For future all continue stable. and be (NN) or current might male #8) personality In the society. (Case male’s multilevel #17, the killed this and show in which #8 selection example case sexual extreme by in an favor observed might have males we tolerance fact, and selection, natural by favored aiosi ..&Tpr ..&Mrhl,TC 20) eiighwtecmue rga CERVUS program computer the how Revising assignment. paternity (2007). in T.C. success Marshall, increases & error genotyping M.L. accommodates Taper, & S.T. Kalinowski, in value MA). adaptive Cambridge, its Press, and reproduction (1997). female S.B. Hrdy, of failure Male-caused ( (1999). marmots W. alpine Arnold, & K. Hacklander, hnpteu roxellana Rhinopithecus nRmaa SuhNepal). (South Ramnagar in ) m .Py.Anthropol. Phys. J. Am. rnsEo.Evol Ecol. Trends amt marmota Marmota h agr fAu eaeadMl taeiso Reproduction of Strategies Male and Female Abu: of Langurs The ). m .Primatol. J. Am. 156 doi:10.1016/j.tree.2020.05.003. . ). 606-613. , ea.Eo.Sociobio. Ecol. Behav. ea.Ecol. Behav. 72 25-32. , 9 10 592-597. , 41 139-150. , tal et Nature 22) utlvlognsto of organisation Multilevel (2020). . Nature o.Ecol. Mol. 303 rnsEo.Evol. Ecol. Trends 381 59-61. , HradUniversity (Harvard . 189-190. , 16 1099-1106. , il Rev. Biol. 18 Presbytis 41–47. , 73 , Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. in,ZF,Fn . hn . i,R,Zag . ag W., Yang, B., Zhang, monkey. R., World Old Liu, free-ranging H., a Chen. in offspring. P., Fan, their Z.F., protect Xiang, females Promiscuous (2004). D.W. 19 MacDonald, personalities. animal & of J.O. evolution Wolff, the favour trade-offs Life-history (2007). 581-584. F.J. Weissing, development. ( & langurs O. troop Hanuman of and history mortality, Life infant (1984). C. Vogel, & H. Loon, (1997). F.B.M. Waal, 136-179. de In: pp. selection. B.) sexual Campbell, langurs and eds Thomas’s investment ( wild Parental in (1972). Vigilance R. (1999). Trivers, J.A.R.A.M. vanHoff, & M. Buul, van ( R.C., Piek, R., Steenbeek, evolutionary and ecological an geladas. syndromes: Primates wild Behavioral in (2004). effect Bruce J.C. ( A Johnson, & (2012). A. J.C. Y., overview. Bell, Beehner, A., & Sih, T.J. Duan, evolu- Bergman, 1222-1225. female A., J., of Lu, E.K., arrest experimental Roberts, Zhang, by triggered matings, adaptation P., male extrapair antagonistic tion. Huang, Sexually increased (1996). W.R. through G., Rice, avoidance Fang, snub-nosed infanticide golden the C.C., facilitate https://doi.org/10.1016/j.anbehav.2019.12.014. of societies dynamics Grueter, Social tilevel X.G., (2009). K. Qi, Watanabe, & W. Ji, 670-679. P.A., Garber, ( B., monkey Li, overview. an X., fertilization: Qi, and mating over conflict Sexual Counter- 361 Female (2006). and A. Strategies G. Male Parker, of Coevolution Conflict: Sexual as strategies. Infanticide (2015). and In: R.A. ecology counterstrategies. Palombit, of female and roles strategies relative Male The Societies Infanticide: (2012) primates: R.A. in Palombit, evolution Social 388-420. (2000). pp. C.P. In: after Schaik, and conflict. van during intersexual loss & Infant C.L. (2019). P.F. Nunn, Fan, & T.Q. gibbons. Bartlett, in O., replacement L.E. societies. male Light, mammalian W.Y., in Brockelman, male C.Y., by Ma, infanticide of monkeys evolution snub-nosed The Sichuan 841-844. (2014). of , E. group Huchard, a & in D. availability Lukas, food China. and Reserve, diet Nature of wild Shennongjia variation of in Seasonal groups (2006). one-male Y.M. within Li, behavior Copulation China. (2007). of Mountains D. Qinling Zhao, & B. Li, rsyi thomasi Presbytis entellus Presbytis 127-134. , 235–259. , Nature 28 hnpteu roxellana Rhinopithecus esMtn,JC,Cl,JP,Kpee,PM,Plmi,RA ik ..,p.432-468. pp. J.B.), Silk, & R.A. Palombit, P.M., Kappeler, J.P., Call, J.C., Mitani, (eds odSrn abPrpc Biol Perspect Harb Spring Cold rnsEo.Evol. Ecol. Trends 163-197. , 381 232–243. , tJdpr(aata/Ida:Rcn bevtosadarcnieaino hypotheses. of reconsideration a and observations Recent India): (Rajasthan/ Jodhpur at ) :teiprac fifniierisk. infanticide of importance the ): natcd yMlsadIsImplications Its and Males by Infanticide ooo h ogte Ape Forgotten The Bonobo: Primates m .Primatol. J. Am. 19 7-7.Sme,V 18) natcd mn rerniglangurs free-ranging among Infanticide (1987). V. Sommer, 372-378. , :fml rnfradoeml ntsuccession. unit one-male and transfer female ): c.Adv. Sci. , 48 m .Primatol. J. Am. oi Primatol Folia 190-196. , 7 a017640. , 81 5 eaav0499. , 23036. e , Uiest fClfri rs,Bree) ike,P. Winkler, Berkeley). Press, California of (University . 10 ea.Eo.Sociobiol. Ecol. Behav. . 68 43 217-233. , rsyi entellus Presbytis -3 of . a or,GS,Leimar, G.S., Doorn, van M., Wolf, 1-23. , tal. et esb a cak ..&Jno,C.H.) Janson, & C.P. Schaik, van by (eds eulSlcinadteDseto Man of Descent the and Selection Sexual 21) otn loaenlnursing allomaternal Routine (2019). .Rpoutv parameters, Reproductive ). 45 hnpteu roxellana Rhinopithecus hls rn.R o.B. Soc. R. Trans. Philos. tal et 137-150. , m .Primatol. J. Am. vlto fPrimate of Evolution 22) Mul- (2020). . rnsEo.Evol. Ecol. Trends nmlBehav Animal Science Nature Science 447 335 71 346 in , , , , Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. ftlrn aeaelne eue hntoeo grsieml fteml oeaeteuwae infant unweaned the tolerate male the if tenures male (b) aggressive unit, than social of kill their longer those not in are than did infant tenures tenures attack unweaned male attack. longer their of (a) without are sooner. and set because male mothers mother first strategies keeping tolerant victims’ the their if of from of if female benefit emigrated offspring of would intervals females next inter-birth male lactating the Therefore, increased as sired obtained offspring. benefits they they the lost aggression, because completely of infants strategy they killing a However, adopted from males success When reproductive of might success. risk detection reproductive less different risk bring to of lead vigilance and can 4 defense Fig size joint OMU female-female efficient. larger stronger more that be because hypothesis also attacked for or evidence killed an being indicating of infanticide death and snub-nosed in infants golden seen of was society 3 multilevel infanticide. increase Fig a male parallel that against No hypothesis counterstrategies for female replacement. evidence facilitate male an monkeys indicating following +infant, rate fetal death and infant fetal seen were increases particular 2 male a of showing Fig died observed females/infants infants circles). of 4 (Number in and shown 2020. survived and are infants 2006 behavior 27 period replacement, study male the during during infanticide present were that infants unweaned 1 Fig captions Figure takeover. male after infant http:// at S1 available Table is (2017). article S.C. this for Alberts, material & Supplementary J. Altmann, E.A., baboons. Archie, MATERIALS male SUPPLEMENTARY L.D., by killing Chow, infant J.B., and Gordon, fetal X., Conditional J.H., Ruan, Grady, W., M.N., Yang, ( Zipple, monkeys H., snub-nosed Yao, grooming golden H., exchange in Chen, monkeys display Y., snub-nosed mouth-opening Female Chen. (2013) Y.J., Li. Zhang, M. & handling. C.C. infant Grueter, and H., sex Yao, for Z., Xiang, in Y., infanticide Yu, Male ( langurs (2016). white-headed Z. wild Xiang in takeovers Zool. & male during M. death Li, infant C.C., and Grueter, H., ( Xu 37 monkey W., snub-nosed Yang, ( golden B., monkey Yang, the H., snub-nosed Yu, Sichuan H., the Yao, primate: polygynous P.A., the Zool. Garber, in Integrat. H., Yao, choice J., Yang, mate Z., and Xiang, A. B., P. Ren, B. Garber, Yang, ( C.C., monkeys snub-nosed Grueter, golden H., in Yao, distributions PeerJ birth X., influencing Qi, potentially W., factors society. Yang, primate multi-level Z.F., P.A. a Xiang, in Garber, males C.C., bachelor Grueter, against H., units Yao, one-male Y., Yu, their B., Yang, Z.F., Xiang, 7-8.Yn . i,T,Wtnb,K,Qn,D,Wn,D a,W 21) aeatcso infants on attacks Male (2013). W. Pan, & D. Wang, D., Qing, K., Watanabe, T., Jin, L., Yin, 175-184. , 8 fe aerpaeet e ae hwdtosrtge,ete grsino oeac,wihwould which tolerance, or aggression either strategies, two showed males new replacement, male After ieecsi ieo oilui oeml nt M)wt n ihu eoddo aeatcson attacks male of recorded without and with OMU) unit, male (one unit social of size in Differences 5 h otn fuwae nat Us of (UIs) infants unweaned of fortune The 365–377. , aitosi ea n natdahrtsdrn h aerpaeet ttsial significant Statistically replacement. male the during rates death infant and fetal in Variations e2892. , hwn aeatcigadfrueo nendifn,otoeo dl eaewt unweaned with female adult of outcome infant, unweaned of fortune and attacking male showing 9 598-612. , lSone PloS hnpteu roxellana Rhinopithecus 8 e74822. , hnpteu roxellana Rhinopithecus 11 rc .Sc B Soc. R. Proc. hnpteu roxellana Rhinopithecus ,asaoal reigprimate. breeding seasonally a ), tal. et tal. et 21)Mjrhsooptblt complex histocompatibility Major (2014) tal et rcyihcsleucocephalusTrachypithecus 284 tal et 21) oilfntoso relaxed of functions Social (2019). 21) ae olcieydefend collectively Males (2014). . , olwn aetkoes f31 of takeovers: male following 20162561. m .Primatol. J. Am. 21) neaiainof examination An (2017). . ). hnpteu roxellana Rhinopithecus hnpteu roxellana Rhinopithecus ol Res. Zool. n.J Primatol. J. Int. 40 76 89-96. , ). 609-617. , Integrat. ). ). Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. natcd uigtesuypro 06ad22.(ubro eae/nat bevdsoigaparticular male a of showing died observed females/infants infants circles). of 4 (Number in and shown 2020. survived and are infants 2006 behavior 27 period replacement, study male the during during infanticide present were that infants unweaned 1 Fig h otn fuwae nat Us of (UIs) infants unweaned of fortune The hnpteu roxellana Rhinopithecus 12 olwn aetkoes f31 of takeovers: male following Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. nrae eese ea et aefloigml elcmn.N aallices a eni et of death snub-nosed in golden seen of was society multilevel infanticide. increase a male parallel that against No hypothesis counterstrategies for female replacement. evidence facilitate male an monkeys indicating following +infant, rate fetal death and infant fetal seen were increases 2 Fig aitosi ea n natdahrtsdrn h aerpaeet ttsial significant Statistically replacement. male the during rates death infant and fetal in Variations 13 Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. nat n natcd niaiga vdnefrhptei htlre M iecnla ols ikof might risk detection less risk to of lead vigilance and can defense size joint OMU female-female efficient. larger more stronger that be because hypothesis also attacked for or evidence killed an being indicating infanticide and infants 3 Fig ieecsi ieo oilui oeml nt M)wt n ihu eoddo aeatcson attacks male of recorded without and with OMU) unit, male (one unit social of size in Differences 14 Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. #18 #17 #16 #15 #14 #13 #12 #11 #10 #9 #8 #7 #6 #5 #4 #3 #2 #1 Events infant unweaned the tolerate S1 male Table the if tenures male (b) aggressive unit, than social of kill their longer those not in are than did infant tenures tenures attack unweaned male attack. longer their of (a) without are sooner. and set because male mothers mother first strategies keeping tolerant victims’ the their if of from of if female benefit emigrated offspring of would intervals females next inter-birth male lactating the Therefore, increased as sired obtained offspring. benefits they they the lost aggression, because completely of infants strategy they killing a However, adopted from males success When reproductive success. reproductive different bring 4. Fig fe aerpaeet e ae hwdtosrtge,ete grsino oeac,wihwould which tolerance, or aggression either strategies, two showed males new replacement, male After aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male 15 DHT NN MH TD GE YH XZ HH4 XB WY NN DY DW DY XX DY DD XY e aeID male New S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male - Nn19 Tt19 Xf19 Ss19 Yb19 Jj2I16 XyI16 NnI15 YbI15 Tt2I15 BbI15 EbI15 XyI14 XeI14 JjI14 Hh2I14 Yy1I13 LnI13 YbI13 LlI13 NnI13 XbI12 Hh3I12 XbI12 - Yy1I09 LnI09 Hh3I08 Hh1I08 QqI08 BbI08 Hh2I08 BbI08 Hh2I08 L1I08 JzI07 JzI07 Jj07 Do eedn offspring dependent of ID S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male - Yes No No No No No No No No No No No No No No Yes No No No No No No No Yes No No No No No No No No Yes Yes Yes No Yes Yes stemte rgatdrn h aevrevent takeover the during pregnant mother the Is S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male - 7 7 6 6 6 6 6 15 15 15 15 6 12 12 12 5 14 14 12 12 12 6 6 6 6 10 10 12 12 12 7 7 5 5 5 6 5 5 M ie(nldn infant) (including size OMU S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male - Yes Yes - No Yes No Yes No No No Yes No No No - Yes No No No No Yes Yes Yes - - Yes Yes No No Yes Yes Yes - - Yes Yes No Yes aeatc nifn rpenn female pregnant or infant on attack Male S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male - Yes(3)a Yes(3)a - Yes(4)a - (2)a Yes - - - (3)a Yes - - - - Yes(3)a - - - - Yes(5)b Yes(3)a Yes(3)a - - Yes(4)c Yes(5)a - - (3)a Yes e (4)b Yes Yes(4)c - - - e (3)a Yes - - eaescoalition* Female’s S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male - rnfrB Transfer A Transfer male Old Transfer e male New Transfer e male New male New male New Transfer e male New e male New male New male Old e male New e male New male New e male New male New Transfer e male New male New male New - Transfer Transfer e male New male New Transfer Transfer Transfer - - - e male New - male New eiec ftemte olwn aereplacement** male following mother the of Residency e male New al S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male - Survive Survive Survive Survive Survive Survive Survive Survive Survive Survive Survive*** Survive Survive Survive Survive Infanticide Survive Survive Survive Survive Survive Infanticide Infanticide - - Survive Survive Survive Survive Survive Survive Survive - - Abortion Infanticide - Abortion aeo h ea ruwae infant unweaned or fetal the of Fate S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male Posted on Authorea 4 Aug 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159657675.51727990 — This a preprint and has not been peer reviewed. Data may be preliminary. h ubro dl eae novdcletv ees r rsne nbakt hc r o nlddterpael taki 1# n 9 a,mastemte steiiitro olciedfne b en h te eae xettemte r h ntao fcletv ees;””masta eiiitro olciedfnei nla.*h ubro dl eae novdcletv ees r rsne nbakt hc r o nlddterpael taki 1# n 9 a,mastemte steiiitro olciedfne b en h te eae xettemte r h ntao fcletv ees;””masta eiiitro olciedfnei nla.*h ubro dl eae novdcletv ees r rsne nbakt hc r o nlddterpael taki 1# n 9 a,mastemte steiiitro olciedfne b en h te eae xettemte r h ntao fcletv ees;””masta eiiitro olciedfnei nla.*h ubro dl eae novdcletv ees r rsne nbakt hc r o nlddterpael taki 1# n 9 a,mastemte steiiitro olciedfne b en h te eae xettemte r h ntao fcletv ees;””masta eiiitro olciedfnei nla.*h ubro dl eae novdcletv ees r rsne nbakt hc r o nlddterpael taki 1# n 9 a,mastemte steiiitro olciedfne b en h te eae xettemte r h ntao fcletv ees;””masta eiiitro olciedfnei nla.*h ubro dl eae novdcletv ees r rsne nbakt hc r o nlddterpael taki 1# n 9 a,mastemte steiiitro olciedfne b en h te eae xettemte r h ntao fcletv ees;””masta eiiitro olciedfnei nla.*h ubro dl eae novdcletv ees r rsne nbakt hc r o nlddterpael taki 1# n 9 a,mastemte steiiitro olciedfne b en h te eae xettemte r h ntao fcletv ees;””masta eiiitro olciedfnei nla.*h ubro dl eae novdcletv ees r rsne nbakt hc r o nlddterpael taki 1# n 9 a,mastemte steiiitro olciedfne b en h te eae xettemte r h ntao fcletv ees;””masta eiiitro olciedfnei nla.*h ubro dl eae novdcletv ees r rsne nbakt hc r o nlddterpael taki 1# n 9 a,mastemte steiiitro olciedfne b en h te eae xettemte r h ntao fcletv ees;””masta eiiitro olciedfnei unclear. is defense collective of initiator he that means ”c” defense; collective of initiator the are mother the except females other the means ”b” defense; collective of initiator the is mother the means ”a”, #9. and #1,#6 in attack repeatedly the included not are which brackets in presented are defense collective involved females adult of number *The unclear. is defense collective of initiator he that means ”c” defense; collective of initiator the are mother the except females other the means ”b” defense; collective of initiator the is mother the means ”a”, #9. and #1,#6 in attack repeatedly the included not are which brackets in presented are defense collective involved females adult of number *The unclear. is defense collective of initiator he that means ”c” defense; collective of initiator the are mother the except females other the means ”b” defense; collective of initiator the is mother the means ”a”, #9. and #1,#6 in attack repeatedly the included not are which brackets in presented are defense collective involved females adult of number *The birth. giving after male old with stay to returned she male, new with stay initially unclear. female is pregant defense the collective B, of male; initiator old he with that stay means to ”c” return defense; and collective male of new initiator with the stay are initially mother infant the unweaned except and females mother other the the A, means unit; ”b” social defense; other collective to of transfer initiator infant the unweaned is and mother mother the the means means ”a”, male;”Transfer” ousted #9. the and means #1,#6 male” in ”Old attack with; repeatedly stay the infant included unweaned not and are mother which the brackets means in male” presented ”New are **, defense collective involved females birth. adult giving of after number male *The old with stay to returned she male, new with stay initially female pregant the B, male; old with stay to return and male new with stay initially infant unweaned and mother the A, unit; social other to transfer infant unweaned and mother the means male;”Transfer” ousted the means male” ”Old with; unclear. stay is infant defense unweaned collective and of mother initiator the he means that male” means ”New ”c” **, defense; collective of initiator birth. the giving are after mother male the old except with stay females to other returned the she means male, ”b” new defense; collective with stay of initially initiator female the pregant is the mother B, the male; means old ”a”, with #9. stay and to #1,#6 return in and attack male repeatedly new the with included stay not initially are infant which unweaned brackets and in mother presented the are A, defense unit; collective social involved other females to adult transfer of infant number unweaned *The and mother the means male;”Transfer” o aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male 16 *,Teifn idwtotml takdrn h winter. the during attack male without died infant The ***, S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male *,Teifn idwtotml takdrn h winter. the during attack male without died infant The ***, S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male *,Teifn idwtotml takdrn h winter. the during attack male without died infant The ***, S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male *,Teifn idwtotml takdrn h winter. the during attack male without died infant The ***, S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male *,Teifn idwtotml takdrn h winter. the during attack male without died infant The ***, S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male *,Teifn idwtotml takdrn h winter. the during attack male without died infant The ***, S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male *,Teifn idwtotml takdrn h winter. the during attack male without died infant The ***, al S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male *,Teifn idwtotml takdrn h winter. the during attack male without died infant The ***, S1 Table aeatcigadfrueo rgatfml,uwae nat ucm fmtes n nendifn olwn aereplacement. male following infant unweaned and mothers, of outcome infant, unweaned female, pregnant of fortune and attacking Male