Female social dynamics in a provisioned free-ranging band of the Sichuan snub-nosed monkey (Rhinopithecus roxellana) in the Qinling Mountains, China

Little information is available on the social dynamics of the Sichuan snub-nosed monkeys (Rhinopithecus roxellana). Here, we provide 5-year observational data on a provisioned free-ranging band in the Qinling Mountains, central China, from October 2001 to December 2005. The results of this study suggested that females place a high priority on maintaining long-term relationships with other female in their one-male units (OMUs). Females had more social interactions with unit members than with nonunit individuals. They showed relatively high rates of affiliation with their unit females, intermediate levels with the resident males, and low levels with individuals outside the unit. Females had few interactions with other resident males. Social interactions, especially affiliative interactions, were more symmetrical between focal females and their unit females than between focal females and the resident males. Thus female social interactions appeared to resemble those of other female-bonded primate societies. On the other hand, female transfers occurred sometimes. We recorded 11 cases of female immigration and 5 cases of female disappearance, 1 case of female intraband shift, and 2 cases of "merging of OMUs," which suggested that unit females might be loose-knit in this species. We discussed influencing factors on female Sichuan snub-nosed monkey social dynamics, by comparing them with those of Hamadryas and gelada baboons that also have multilevel societies.     

Social organization of black-and-white snub-nosed monkeys (Rhinopithecus bieti) at Deqin, China

Data on social organization of two bands of black-and-white snub-nosed monkeys (Rhinopithecus bieti) were collected when the monkeys were crossing an open spot at Nanren and Bamei (northwest of Yunnan, China) using a sampling rule where individuals within one social unit are spatially closer to each other than individuals between social units. The typical pattern of social organization in this sample was multiple adult females (AFs) and their offspring with one adult male (AM) in a one-male unit (OMU), similar to that of many other colobines. In such units, on average one male is associated with 4.0 AFs and 2.5 of their offspring. Moreover, there are multimale/multifemale units and monogamous units besides OMUs. All bisexual units traveled together with at least one all-male unit as a cohesive band. In two bands of monkeys, 87% of AMs in bisexual units were within OMUs, 7.8% within monogamous units and 5.2% within multimale, multifemale units. In the Bamei band, 6.7% of AMs were in the all-male unit. The size of OMUs in the Nanren band was larger than that of the Bamei band, with more AFs and juveniles, which may be related to better conservation in the Nanren band's habitat. For the Nanren band, the average number of AFs in OMUs varied across time, increasing from 4.3 in 1994 to 5.1 in 2001, and then decreasing to 3.8 in 2005. This article suggests three possible explanations for this variation, but more data are needed for these hypotheses to be tested.     

Impact of male takeover on intra-unit sexual interactions and subsequent interbirth interval in wild Rhinopithecus roxellana

Data were collected on sexual interactions before and after a male takeover of a one-male unit (OMU) of Sichuan snub-nosed monkeys (Rhinopithecus roxellana) in the Qinling Mountains, China. The original unit consisted of an adult male, 2 adult and 2 subadult females, 2 female juveniles and a single infant. Following the takeover, the new resident male copulated with 1 adult female, which was not lactating. Subsequent to the disappearance of her infant, the second (lactating female) entered breeding condition and began to solicit copulation with the new resident male. The subadult females also engaged in matings with the new male. The new resident male was observed mating, on 3 occasions, with females in 2 other OMUs. These are the first observations of sexual behaviour in free-ranging Sichuan snub-nosed monkeys after an OMU takeover. Sexual interactions play an important role in establishing relationships between the new male and resident females in the OMU.     

A Proximity-Based Social Network of a Group of Sichuan Snub-Nosed Monkeys (Rhinopithecus roxellana)

Studies of spatial proximity between individuals are important for an understanding of social structure because animals are more likely to interact with individuals in close spatial proximity. Here, we apply social network analysis to proximity data collected between 2001 and 2003 from an individually identified, provisioned, free-ranging band of Sichuan snub-nosed monkeys (Rhinopithecus roxellana) in the Qinling Mountains, central China. We aimed to quantify the social network structure and to gain insight into each individual’s position within the social network. The overall network connectivity of the study band was sparse, with a low group density of 0.17. We identified nine one-male-multifemale units (OMUs) in the study band using hierarchical cluster analysis, which confirms that this species forms a multilevel society in its natural habitat. Based on sex differences in eigenvector and betweenness centralities, adult females have more important social roles than males. Among females, lactating females scored higher betweenness and eigenvector centralities than other females. However, our results do not suggest the existence of key individual(s) in the social network of the study band. The global clustering coefficient characteristic of the band was 0.3?±?0.1, with little variation between individuals, suggesting that the removal or death of any specific individual would not significantly disrupt its general network structure. Our results also show that proximity commonly occurs among unit members, but can also occur between females of different OMUs. These observations suggest that snub-nosed monkeys have a loose-knit or fluid rather than a rigid female-bonded social system, which may be a common trend for species living in multilevel societies.     

Female choice impacts resident male takeover in golden snub-nosed monkeys(Rhinopithecus roxellana)

In primate species with social systems consisting of one-male breeding units(OMUs), resident male takeover represents a major challenge to individual reproductive success and mating strategies. The golden snub-nosed monkey(Rhinopithecus roxellana)is characterized by large multilevel societies(MLS)comprised of several OMUs and all-male units(AMUs);however, the factors and mechanisms associated with resident male takeover, which offer important insight into primate reproduction and social strategies, are still poorly understood. Based on 5-year monitoring data from a free-ranging herd of golden monkeys from the Qinling Mountains in China, we categorized three phases of an OMU, that is, a rising phase,developing phase, and declining phase. The rising and declining phases were unstable periods in which male takeover in an OMU might occur. Factors causing takeover, such as leader male rank, fighting ability, reproduction rate, and affiliation(proximity,allogrooming), were analyzed for males and females and for different OMUs. Results indicated that the new resident male's fighting ability was lower than that of the former resident male in 23 cases. After replacement, the rank order of the new resident male significantly declined. Females involved in a takeover increased their distance from the resident male and decreased mating frequency during the three months prior to takeover. Females with infants under one-year-old had a marked effect on the specific time of takeover occurrence. These results suggested that female choice was the main factor deciding whether a takeover attempt was successful. Furthermore, rather than male conflict, females more often initiated and affected takeover and outcome, implying that the social status and competitive ability of the males played lesser roles during takeover.     

The role of kinship in the formation of a primate multilevel society

A small number of mammalian species live in a modular or multilevel society in which several individual social/reproductive units called one-male units (OMUs) are embedded within a large cohesive band. Factors that affect band composition and stability are poorly understood. In this study we examined the role of kinship in the formation and maintenance of a multilevel society in an endangered population of golden snub-nosed monkeys (Rhinopithecus roxellana). From 2005 to 2011, we obtained genetic samples from 86 individuals (including 88.9% of leader males and 80.5% of adult females) living in a band of 8–10 OMUs. We used microsatellite genotyping to identify patterns of relatedness and individual transfer. We found that adult females residing in the same OMU were more closely related to each other than to a random set of females drawn from the band and that females tended to disperse into OMUs that contained female relatives. In addition, adult females who transferred were not more closely related to their previous leader male than to the leader male of their new OMU. These results support the contention that kin bonds contribute importantly to the formation and stability of this primate multilevel society by influencing a female's decision to remain in her current OMU, or during transfer, which new OMU to enter. Am J Phys Anthropol 156:606–613, 2015. © 2014 Wiley Periodicals, Inc.     

Female preferences for male golden snub-nosed monkeys vary with male age and social context

Age is a key factor affecting sexual selection, as many physical and social traits are age-related. Although studies of primate mate choice often consider particular age-related traits, few consider the collective effects of male age. We tested the hypothesis that female golden snub-nosed monkeys Rhinopithecus roxellana prefer prime aged males (10–15 years) over younger and older males. We examined a habituated, provisioned troop during a 3-year study in the Qinling Mountains, China. Prime age males were more likely to be resident males of 1-male units (OMUs) than males of other ages. Since females are free to transfer between OMUs, the number of females per OMU can be indicative of female preferences. We examined the number of females per OMU, and found that it increased with resident male age up to 7–8 years, and declined after 12 years, such that prime age resident males had more females than other resident males. Females also initiated extra-unit copulations with high-ranking prime age males at significantly higher rates than with other males. Nevertheless, females tended to transfer from OMUs with high-ranking, older resident males to those with low-ranking, younger resident males. Thus, females appear to use different strategies when choosing social mates and extra-unit mates (i.e., different social contexts). We speculate that females may perceive early signs of aging in males and trade off the benefits and costs of high rank versus male senescence. This study lays the groundwork for future studies that examine possible direct and indirect benefits of such strategies.     

A preliminary study of food transfer in Sichuan snub-nosed monkeys (Rhinopithecus roxellana)

Food transfer happens regularly in a few nonhuman primates species that are also characterized by remarkable social tolerance. Sichuan snub-nosed monkeys (Rhinopithecus roxellana), or golden monkeys, which exhibit high social tolerance in their social relationships are thus of interest to see whether tolerance would extend to food transfer. In this study, branch feeding activity was observed in a semi-captive group of Sichuan snub-nosed monkeys, which consisted of 10 subjects that included a one-male unit (OMU) and an all-male unit (AMU). We recorded 1,275 food interactions over 27 days, and 892 instances of food transfer. The most commonly observed types of food transfer behavior were co-feeding (62.1%) and relaxed claim (22.8%). Of 892 food transfers, 756 (84.8%) took place in the OMU, most of which were among adults (34.7%) and among juveniles (42.1%). The transfer success rate was high in both the cases (87.9% for adults and 78.9% for juveniles). Food transfer in the AMU took place less often than that among adults in the OMU though with similar high transfer success. Food transfer between the OMU and AMU was limited to juvenile males from the OMU and adults from the AMU. These results provide the first evidence of food transfer in golden monkeys and suggest that tolerant social relationships in golden monkeys make transfer possible.     

Social dynamics of the golden snub‐nosed monkey (Rhinopithecus roxellana): female transfer and one‐male unit succession

Among primates that form multilevel societies, understanding factors and mechanisms associated with the movement of individuals between groups, clans, and one‐male social units offers important insight into primate reproductive and social strategies. In this research we present data based on an 8‐year field study of a multilevel troop of Sichuan snub‐nosed monkeys (Rhinopithecus roxellana) in the Qinling Mountains of China. Our study troop contained 78–126 individuals, and was usually organized into 6–8 one‐male units (OMU). The majority of OMUs were composed of networks of unrelated females and their offspring. We found that 59.7% (43/72) of subadult and adult females in our study troop transferred between OMUs (n=66) or disappeared (n=7) from the troop. In the majority of cases, two or more females transferred together into new OMUs or troops. In R. roxellana, new OMUs formed in several ways. During 2001–2008, 16 adult males appeared in the study troop. Over this period, we observed 13 different males who became harem leaders either by taking over an existing harem or by attracting females from other OMUs into their harem. We also observed four OMUs from a neighboring troop to successfully immigrate into the study troop. The number of individuals in these newly immigrated OMUs was significantly smaller than that number of individuals in resident OMUs. During harem formation, fighting between adult males was rarely observed, and female mate choice appeared to play a crucial role in harem male recruitment and replacement. These results suggest that golden snub‐nosed monkeys are organized in a nonmatrilineal social system. Female mate choice and possibly incest avoidance appear to play important roles in female transfer, male tenure, and OMU stability. Am. J. Primatol. 71:670–679, 2009. © 2009 Wiley‐Liss, Inc.     

利用过夜地粪便估计白马雪山黑白仰鼻猴种群大小和组成

介绍了应用过夜地粪便来估计白马雪山黑白仰鼻猴群大小和组成的一种方法。该物种以单雄多雌单元和全雄组的形式在树上过夜。粪粒根据其大小可分为3种类型成年雄性的(最大)、成年雌性的(中等大小)和未成年个体的(最小)。2000—2001年,搜集了滇西北白马雪山国家级自然保护区北部南任村(99°04′E,28°34′N)附近黑白仰鼻猴群每个季节2个过夜地的粪粒。根据2001年11月猴群通过开阔地的数据来确定猴群组成。每个季节,由于单雄多雌单元的成年个体数与其粪粒数正相关,所以二者回归直线的斜率可以看作是每个个体每晚的平均排便量。由于该物种的栖息地主要为高山峡谷,而且能见度较低,因此,利用过夜地粪便比以前通过猴群活动痕迹来估计猴群大小和组成相对准确、可靠。从估计成年雌性个体数的角度看,利用粪粒来估计种群大约有9·4%的偏差。导致偏差的可能原因有杂草和灌丛对粪粒准确计数的影响、个体排粪率的差异以及成年雄性最小粪粒与成年雌性最大粪粒的混淆等。该方法适应于栖息地和主要食物与本文研究种群相似的其他种群。     

Social functions of relaxed open-mouth display in golden snub-nosed monkeys(Rhinopithecus roxellana)

Relaxed open-mouth display serves important social functions in relation to submission, reconciliation affiliation and reassurance among non-human primate societies; however, quantitative evidence on this behavior remains insufficient among multi-level social groups. From July to November 2016, we examined four potential functions of the relaxed open-mouth display during pairwise, intra-unit social interactions among 18 free-ranging adult and sub-adult golden snub-nosed monkeys(Rhinopithecus roxellana) who belonged to three one-male, multi-female units(OMU) at Dalongtan, Shennongjia National Park China. Results showed that: compared with no relaxed open-mouth display,(1) the occurrence of displacement by a dominant individual approaching a subordinate was lower and the distance of the subordinate to the approaching dominant was shorter when the subordinate showed open-mouth display;(2) relaxed open-mouth display reduced the probability of continued attack for victims of aggression and allowed victims to achieve closer proximity to the aggressor during post-conflict periods;(3) relaxed open-mouth display by dominant individuals allowed them to achieve closer proximity to subordinates; and(4) the exchange of relaxed open-mouth display had a greater impact on the outcome of interactions than one individual alone giving this signal. These findings suggest that relaxed open-mouth display serves important functions regarding submission, reconciliation, affiliation and reassurance in coordinating social interactions within OMUs in golden snub-nosed monkeys.     

Anogenital distance as a measure of male competitive ability in Rwenzori Angolan colobus

Anogenital distance (AGD) is positively correlated to fetal androgen exposure and developmental masculinization in mammals. Independent of overall body size, AGD shows a strong positive correlation with male fertility and in rodents, AGD is a good indicator of male competitive ability and is associated with female choice. We hypothesized that AGD will also predict male competitive ability in non-human primates. To test this, we measured AGD noninvasively with a parallel laser in a wild population of Angolan colobus monkeys (Colobus angolensis ruwenzorii) in Uganda and correlated to it to their social structure. C. angolensis ruwenzorii form a multilevel society with both one-male/multifemale units (OMUs) and multimale/multifemale units (MMUs). We compared AGD in males from five OMUs and six MMUs and related it to their fecal androgen metabolite concentrations, dominance rank and body size, and to the number of females in their unit. Males in OMUs had greater access to females, so were predicted to have longer AGDs, but this was not found. AGD also did not correlate overall with mean fecal androgen metabolites in MMUs. However, AGD was correlated with dominance rank in MMUs, demonstrating that higher-ranking males in these multimale units had longer AGDs. Body size did not show the same relationship with dominance rank, suggesting that male rank was not just a reflection of absolute male size. Our findings indicate that AGD predicts male competitive ability in this species and that it may be a useful correlate throughout the non-human primates. These results also support the idea that prenatal androgen exposure increases the likelihood of the expression of behaviors that maintain high dominance rank.     

九寨沟自然保护区川金丝猴的分布及种群数量

2014年2月至5月,对四川省九寨沟自然保护区野生川金丝猴的种群分布、数量及其社会结构进行调查,初步探讨该种群与其他种群在外部形态和社会结构方面的差异性。采用"V"型路线法调查,发现九寨沟保护区共有川金丝猴4群。然后结合长时间的持续跟踪观察,证实家庭单元19个,全雄群3个,能区分不同个体296只。OMU大小平均为12.7只,其中OMU内的成年个体占38%,青少年个体占53%,幼仔占9%。成年与青少年比例为1∶1.39,成年雌雄的比例为3.84∶1。与秦岭种群的外部形态相比,该种群的成年雄性更为壮大,毛发弯曲程度明显;成年雌性毛发略显灰黑色,金黄色暗淡,大部分个体嘴角长有很小的肉瘤;青少年个体毛色泛白。亦发现九寨沟种群倾向于选择海拔偏高(2400—3550 m)的针叶林带活动。     

雄性川金丝猴对雌性的受孕认知与等级识别

社会认知是群居动物适应复杂环境和社会生活能力的体现,致力于理解自身、他人与社会。认知有利于促进利益分配的优化,而行为认知策略则是关注的热点问题。但迄今为止,川金丝猴如何进行社会地位与受孕状态等基础认知尚不清楚。本研究通过投食招引,在个体识别和等级判定的基础上,以栖息于秦岭观音山自然保护区熊猫谷景区的一个半野生川金丝猴群为对象,采用瞬时扫描和焦点动物取样法,观察并收集了猴群中6个一雄多雌单元成年雌雄个体的空间位置、邀配、交配及产仔行为数据,结合食物密度梯度"同心圆"和受孕周期,推测雌性成功受孕的交配时间段,运用配对t-检验统计证实受孕前后雌性的交配实现率差异显著(t=4.527,P=0.001),表现为受孕前雌性邀配行为获取的雄性响应率明显高于受孕后;通过Spearman相关性检验发现,雌性受孕前的交配实现率(R=0.527,P=0.006)及受孕时间(R=0.556,P=0.049)与自身等级均显著正相关,即高等级雌性邀配行为的雄性响应率及与雄性生殖性交配的时间明显高于或早于低等级雌性。结果表明,雄性川金丝猴拥有认知单元内成年雌性受孕状况和等级地位的能力,这种认知能力影响着雄性的性行为方式。     

中国云南塔城滇金丝猴的社会结构

在云南塔城附近的山地森林中,我们利用摄象机记录了一群滇金丝猴通过山谷和在水源地饮水的过程。我们利用这些记录分析这群滇金丝猴的种群大小、组成成员以及个体的空间分布,并利用连续通过个体之间的时间间隔来揭示社会结构。塔城滇金丝猴群由366个个体组成,是目前所知最大的滇金丝猴群体。猴群内雌性个体较多。猴群可以分为26个单位,其中有19个单雄多雌单位、5个多雄多雌单位和2个全雄单位。统计结果表明单位内个体间的时间间隔显著比单位间的短。2/3以上的亚成年雄性个体和全雄单位一起活动。猴群在地面活动时全雄单位位于猴群的外围,充当前卫和后卫。塔城猴群的生命表组成结构与吾牙普亚猴群的大致一样,同样是多层社会结构,单雄多雌和全雄单位构成猴群的主体。亚成年雄性多数与全雄单位一起活动,表明雄性可能会从其出生的单雄多雌单位中迁移出来。     

Food Transfer in Sichuan Snub-nosed Monkeys (<Emphasis Type="Italic">Rhinopithecus roxellana</Emphasis>)

Food transfer behavior provides a way to distribute food resources among individuals. It is not confined to kin, but also occurs among genetically unrelated individuals. Food transfer among nonkin may result from byproduct mutualism, reciprocal altruism (RA), or tolerated scrounging (TS). Sichuan snub-nosed monkeys (Rhinopithecus roxellana) exhibit a high level of social tolerance, and researchers have observed food transfer behavior in the wild. However, little is known about how tolerant social relations influence food transfer in this species. We recorded food-related interactions and social behavior in a group of captive Sichuan snub-nosed monkeys. Our findings suggest that the monkeys develop partner preference in food transfer behaviors. Moreover, individuals rely primarily on nonharassed approaches to claim food, suggesting that the TS model alone cannot explain their food transfer. Food transfer in this species may be a form of mutualism, in which an individual benefits on an immediate basis by fostering a preferred and tolerant relationship. However, we cannot rule out the possibility of reciprocal altruism. Future studies should record the temporal delay of social exchange to distinguish between mutualism and reciprocal altruism.     

One-male units and clans in a colony of hamadryas baboons (Papio hamadryas hamadryas): effect of male number and clan cohesion on feeding success

In the multilevel societies of hamadryas baboons, adult males can be attached to single one-male units (OMUs) or to clans containing several such OMUs. This paper examines the effect of male number and rivalry between males within a clan on their ability to compete for access to a clumped food resource. The data come from a study of a multilevel colony of hamadryas baboons (Papio hamadryas hamadryas) housed at the Madrid Zoo. The colony consisted of 12 harem-holding males and 40 sexually mature females, and was organized into five single OMUs and two clans (containing three and four OMUs, respectively). The top-ranking male of one of the clans was removed and later reintroduced, so the study involved an analysis of the composition of clans and OMUs and of the males' use of the feeding area across three study periods: preseparation, separation, and reintroduction. The findings reported indicate that both males and females derived clear advantages in the context of contest competition for access to clumped food if they were members of clans, because the males and females from large clans had a feeding advantage over those from smaller clans and single OMUs. Furthermore, rivalry among males within the clan reduced their ability to compete for food against males outside their clan. This paper provides empirical evidence for one of the potential advantages that hamadryas males may enjoy if they are attached to clans, and also provides empirical support for the general hypothesis that a large number of males in a group may provide fitness-related benefits to the group members, provided they are able to cooperate with each other.     

秦岭川金丝猴成年雌性优势资源占有与社会等级的判定

群体生活利弊并存,有利的一面是降低了被捕食的风险和提高了优势资源共享度,不利的一面是时刻面临群内资源竞争。等级关系是群居非人灵长类物种中最常见的一种资源调节方式,制约着优势资源在个体间的分配。为了探讨川金丝猴群的优势资源占有规律与社会等级的关系,本研究以秦岭北坡中段周至国家级自然保护区内玉皇庙村的一个人工投食川金丝猴的4 个一雄多雌单元(OMU)内成年雌性为研究对象,从2009 年9 月至2010 年1 月通过设计同心圆,以优势资源区域占有频次、时间等参数指标计算获取程度,从而对该猴群成年雌性的等级关系进行判定。结果证实取食和休息阶段的等级顺序基本上一致,通过Spearman 相关性非参数检验将投食和休息时的等级顺序与早期采用取代行为判定结果进行比较,具有极显著的正相关性。本研究表明利用优势资源占有确立单元内雌性等级具有一定的可行性,并且在野外操作和数据统计上更简单易行,可以在更短的时间内迅速判定雌性等级,或许能为野外投食和笼养条件下非人灵长类雌性等级的研究提供一种新的判定方法。     

The fourth level of social structure in a multi‐level society: ecological and social functions of clans in hamadryas baboons

Hamadryas baboons are known for their complex, multi‐level social structure consisting of troops, bands, and one‐male units (OMUs) [Kummer, 1968. Social organization of hamadryas baboons. Chicago: The University of Chicago Press. 189p]. Abegglen [1984. On socialization in hamadryas baboons: a field study. Lewisburg, PA: Bucknell University Press. 207p.] observed a fourth level of social structure comprising several OMUs that rested near one another on sleeping cliffs, traveled most closely together during daily foraging, and sometimes traveled as subgroups independently from the rest of the band. Abegglen called these associations “clans” and suggested that they consisted of related males. Here we confirm the existence of clans in a second wild hamadryas population, a band of about 200 baboons at the Filoha site in lowland Ethiopia. During all‐day follows from December 1997 through September 1998 and March 2005 through February 2006, data were collected on activity patterns, social interactions, nearest neighbors, band fissions, and takeovers. Association indices were computed for each dyad of leader males, and results of cluster analyses indicated that in each of the two observation periods this band comprised two large clans ranging in size from 7 to 13 OMUs. All band fissions occurred along clan lines, and most takeovers involved the transfer of females within the same clan. Our results support the notion that clans provide an additional level of flexibility to deal with the sparse distribution of resources in hamadryas habitats. The large clan sizes at Filoha may simply be the largest size that the band can split into and still obtain enough food during periods of food scarcity. Our results also suggest that both male and female relationships play a role in the social cohesion of clans and that males exchange females within clans but not between them. Am. J. Primatol. 71:948–955, 2009. © 2009 Wiley‐Liss, Inc.     

Alternative routes to the leader male role in a multi-level society: follower vs. solitary male strategies and outcomes in hamadryas baboons

The nested one-male units (OMUs) of the hamadryas baboon are part of a complex social system in which "leader" males achieve near exclusive mating access by forcibly herding females into permanent consortships. Within this multi-level social system (troops, bands, clans and OMUs) are two types of prereproductive males--the follower and solitary male--whose different trajectories converge on the leader role. Here we compare OMU formation strategies of followers, who associate with a particular OMU and may have social access to females, with those of solitary males, who move freely within the band and do not associate regularly with OMUs. Data were derived from 42 OMU formations (16 by followers and 26 by solitary males) occurring over 8 years in a hamadryas baboon band at the Filoha site in Ethiopia. "Initial units" (IUs) with sexually immature females (IU strategy) were formed by 44% of followers and 46% of solitary males. The remaining followers took over mature females when their leader was deposed (challenge strategy) or disappeared (opportunistic strategy), or via a seemingly peaceful transfer (inheritance strategy). Solitary males took over mature females from other clans and bands, but mainly from old, injured or vanished leaders within their clan (via both the challenge and opportunistic strategies). Former followers of an OMU were more successful at taking over females from those OMUs than any other category of male. Despite this advantage enjoyed by ex-follower leaders, ex-solitary leaders were equally capable of increasing their OMU size at a comparable rate in their first 2 years as a leader. These results demonstrate the potential for males to employ both multiple roles (follower vs. solitary male) and multiple routes (IU, inheritance, challenge, opportunistic) to acquire females and become a leader male in a mating system characterized by female defense polygyny in a competitive arena.