Female "dispersal" in hamadryas baboons: transfer among social units in a multilevel society

Unlike most cercopithecines, hamadryas baboons (Papio hamadryas hamadryas) are characterized by female-biased dispersal. To clarify this pattern within the context of their hierarchical social system (comprising one-male units, clans, bands, and troops), we report here 7 years of data on female transfers among social units in wild hamadryas baboons in Ethiopia. Female tenure in one-male units (OMUs) ranged from 1 to 2,556 days (N = 208) and survival analysis revealed a median tenure length of 1,217 days (40 months). Changes in OMU membership consisted almost exclusively of takeovers by males, not voluntary transfer. Of 130 takeovers, 67% occurred within the band and 33% across bands, and, of the 22 takeovers for which we have clan membership data, 77% occurred within, not between, clans. These results reinforce the notion that hamadryas female dispersal is not analogous to sex-biased dispersal in other taxa, because (1) at least in Ethiopian populations, females do not disperse voluntarily but are transferred, often forcibly, by males; (2) only dispersal between bands will promote gene flow, whereas females are most often rearranged within bands; (3) hamadryas females undergo social dispersal but not usually locational dispersal; and (4) while male hamadryas are far more philopatric than females, they have been observed to disperse. It thus appears that the ancestral baboon pattern of female philopatry and male dispersal has evolved into a system in which neither sex is motivated to disperse, but females are forcibly transferred by males, leading to female-mediated gene flow, and males more rarely disperse to find females.     

Migration of male hamadryas baboons into anubis groups in the Awash National Park,Ethiopia

Among “savanna” baboons, males are the dispersing sex, and females are philopatric. Despite clear evidence for migration of adult males at Erer-Gota, Ethiopia (Abegglen, 1984), it is generally believed that a different pattern-dispersal only by female transfer-is found in hamadryas baboons,Papio hamadryas hamadryas (Pusey and Packer, 1987; Pusey, 1988; Stammbach, 1987). Since the late 1960's, there have been isolated observations of hamadryas males migrating into anubis groups in the Awash National Park, Ethiopia (Nagel, 1973; Kawai and Sugawara, 1976; Sugawara, 1982). Since 1983, we have observed 11 individually identified adult hamadryas immigrants in four anubis groups above the Awash Falls and have trapped and tagged 9 of them. One subadult male was also captured and marked. Repeated visits to the study site allow us to document long-term residence of these “cross-migrant” males in their host groups. The longest-resident male has been in the same group for 5 years or more; a conservative estimate of the average length of residence is 3 years. We estimate that 25 hamadryas males have moved into this ozne over the last 15 years. Although larger than the hamadryas males captured in 1973, all but one of our cross-migrants appear phenotypically hamadryas. By comparing, the ages of our cross-migrants with Abegglen's account of the typical hamadryas male life-history, we have found that the adult hamadryas males seem to immigrate at ages consistent with having left their     

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.     

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.     

Early sexual maturity in male hamadryas baboons (Papio hamadryas hamadryas) and its reproductive implications

We present data on sexual maturity in young hamadryas baboon males (Papio hamadryas hamadryas) and its reproductive consequences in a large captive baboon colony. Hamadryas baboons live in a multilevel social system, with one-male units (OMUs) as the smallest social entity. Male leaders of OMUs are believed to monopolize matings within their OMUs; hence mating is believed to be polygynous and monandrous. In a captive colony of hamadryas baboons, we found evidence that young males less than 4 years old fathered at least 2.5% of 121 offspring born subsequent to vasectomy of all adult males, and males aged 4-5 years fathered at least 16.5% of the offspring. Additional evidence that these young males are able to sire offspring came from a morphological comparison of sperm from hamadryas males of different ages. The sperm of a 48-month-old hamadryas baboon were morphologically indistinguishable from viable sperm from adult males, whereas sperm from a 45-month-old male showed some aberrations. If successful copulations by adolescent males constitute a regular pattern even in free-ranging hamadryas baboons, a hamadryas male's chances to reproduce would not be limited to his role as an OMU leader as previously assumed, and a male's reproductive career would consist of two phases: the adolescent phase, and the OMU leader male phase.     

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.     

The Socioecology of Network Scaling Ratios in the Multilevel Society of Hamadryas Baboons (Papio hamadryas hamadryas)

Multilevel or modular societies characterize a range of mammalian taxa, allowing social groups to fission and fuse in response to ecological factors. The modular society of hamadryas baboons has previously been shown to consist of 4 levels: troop, band, clan, and one-male unit (OMU). A recent study by Hill et al. (Biology Letters 4:748–751, 2008) revealed a mean scaling ratio across successive levels of multilevel societies of ca. 3; this was consistent across elephants, orca, geladas, and hamadryas baboons. Here we reanalyze the scaling ratio for hamadryas baboons with previously unavailable data from Filoha. Our analysis revealed a mean scaling ratio for hamadryas of 3.28 without data on the hamadryas clan layer of organization at Filoha, but a ratio of 6.17 with these data included. This discrepancy is due to the large clan and band sizes at Filoha yielding a larger than average gap between the OMU and the clan. Further analysis revealed subsets of OMUs within clans, suggesting a 5th level of society in this population. When this 5th layer of social structure is included in the analysis, the scaling ratio at Filoha is consistent with that of other hamadryas populations and other taxa. These results suggest that a consistent mammalian scaling ratio can be used to detect previously hidden levels of organization within societies and to predict their sizes in taxa for which detailed behavioral data are not available.     

One-male groups and intergroup interactions of mountain baboons

The behavioral ecology of mountain baboons, Papio ursinus,cuts across the traditional dichotomy between savannah and desert baboons, providing fresh opportunities to examine socioecological relationships. Mountain baboons were studied at a site where covariation in altitude and group size helps to clarify the influence of ecology on social behavior. One-male groups and lone males were regularly found in the highest-altitude zones. In encounters with multimale groups, a one-male group retreated; the group’s single male attacked his females, herding them far away from the multimale group. The male himself then approached and watched (and often loud-called at) this group. In a two-male group, the younger male herded the females, while the older sometimes defended the group. One-male groups did not show regular, stable aggregations, but two such groups seasonally coordinated their ranging “in tandem.” Ecological explanations for these behaviors and their relationship to the behavior of hamadryas baboons are evaluated, and a new theory of the origin of one-male groups in baboons is developed.     

Cerebrospinal Fluid Monoaminergic Metabolites in Wild Papio anubis and P. hamadryas are Concordant with Taxon-specific Behavioral Ontogeny

We used a cross-sectional sample to compare ontogenetic trajectories in the concentrations of monoamine neurotransmitter metabolites in cerebrospinal fluid of wild anubis (Papio anubis, n = 49) and hamadryas (P. hamadryas, n = 54) baboons to test the prediction that they would differ, especially in males, in association with their distinct behavioral ontogenies. Values of all 3 metabolites [3-methoxy-4-hydroxyphenylglycol (MHPG), the norepinephrine metabolite; 5-hydroxyindoleacetic acid (5-HIAA), the serotonin metabolite; and homovanillic acid (HVA), the dopamine metabolite] declined consistently with dentally-calibrated maturation, and few taxon-related differences were apparent among juveniles. Adult females were too few for adequate comparison, but a discriminant function suggested that they might differ by taxon. Adult males of the 2 species differed strikingly from juveniles and from each other. Contrary to our initial hypothesis, adult male anubis had significantly lower HVA and MHPG, and higher 5-HIAA levels, than predicted from the overall, age-related trend, and MHPG continued to decline with age among adults. As young adults, male hamadryas had low 5-HIAA and a high HVA/5-HIAA ratio, while older males [normatively one-male unit (OMU) leaders] showed a reversal in the trend, with 5-HIAA rising and the HVA/5-HIAA ratio tending to fall. We speculate that the results are related to the dispersing and philopatric ontogenies of anubis and hamadryas males, respectively. Adult male anubis, whose fitness depends on building social networks with nonkin, have high relative serotonin activity, commonly associated with greater social circumspection and skill. Young adult male hamadryas, living among agnatic kin and mating opportunistically, exhibit low 5-HIAA levels, generally associated with impulsivity and social irresponsibility. This reverses as a male approaches the age at which he is normatively the leader of a one-male unit (OMU), and his fitness depends on his maintaining stable relationships with other leaders and with females. An erratum to this article can be found at     

Social System of a Hybrid Baboon Group (Papio anubis × P. hamadryas)

We describe the social organization, mating system, and social structure of a group of hybrid baboons (Papio anubis×P. hamadryas) in Ethiopia's Awash National Park. The group contained elements of both hamadryas and anubis societies. Overall, the group was a multimale, multifemale group that lacked cohesion and frequently formed subgroups. Subgroup formation was more strongly associated with predation risk than food availability. Although there were several hamadryas-like one-male units OMUs within the group, there was no evidence of a hamadryas multilevel society. Male and female members of OMUs were phenotypically more hamadryas-like than non-OMU individuals. The group contained substantial variation in the strength of inter- and intrasexual bonds: some females primarily groomed males while other females primarily groomed females, and the patterns were consistent with the OMU substructure. Despite some promiscuous mating, mating was biased towards the hamadryas condition for all group members. Additionally, rates of immigration and emigration were very low, and mean pairwise relatedness within the group is rising. For measures of intersexual bonding, all members of the group were intermediate between anubis and hamadryas individuals in less hybridized groups. The group was phenotypically and behaviorally more intermediate than it was in the 1970s (Sugawara, K. (1988). Primates 29: 429–448.) and the changes may indicate a relatively young and dynamic hybrid zone.     

Life history of hamadryas baboons: Physical development,infant mortality,reproductive parameters and family relationships

Demographic and life history parameters were estimated for a band of free-ranging hamadryas baboons, observed for 5.5 years in Ethiopia. Age-related changes in body weight and dentition were found to be delayed relative to laboratory-reared baboons. On the average, females reached menarche at 4.3 years of age and had their first infant at the age of 6.1 years. The mean interbirth interval was 24 months if the infant survived this period. The survival of infants and juveniles was higher compared to Amboseli yellow baboons, but somewhat lower than in gelada baboons in the Simen Mountains. Males acquired their first juvenile or adult female at the age of 8.5 to 11 years. Male-female pair-bonds lasted several years in most cases. The Cone Rock baboons were organized in a four-level social structure. The troop could split into bands, bands were divided into clans, and clans into one-male units with bachelor followers. The exchange of individuals between social units predominantly occurred within the band. All males of known origin became adult members of their presumed natal clan. Most females transferred also within clans, and juvenile females tended to remain in their natal clan. Females lost by one male to several rivals tended to reassemble in the same new one-male units later on.     

Copulation patterns in captive hamadryas baboons: a quantitative analysis

For primates, as for many other vertebrates, copulation which results in ejaculation is a prerequisite for reproduction. The probability of ejaculation is affected by various physiological and social factors, for example reproductive state of male and female and operational sex-ratio. In this paper, we present quantitative and qualitative data on patterns of sexual behaviour in a captive group of hamadryas baboons (Papio hamadryas), a species with a polygynous–monandric mating system. We observed more than 700 copulations and analysed factors that can affect the probability of ejaculation. Multilevel logistic regression analysis and Akaike’s information criterion (AIC) model selection procedures revealed that the probability of successful copulation increased as the size of female sexual swellings increased, indicating increased probability of ovulation, and as the number of females per one-male unit (OMU) decreased. In contrast, occurrence of female copulation calls, sex of the copulation initiator, and previous male aggression toward females did not affect the probability of ejaculation. Synchrony of oestrus cycles also had no effect (most likely because the sample size was too small). We also observed 29 extra-group copulations by two non-adult males. Our results indicate that male hamadryas baboons copulated more successfully around the time of ovulation and that males in large OMUs with many females may be confronted by time or energy-allocation problems.     

Longitudinal changes of dominance rank among the females of the Koshima group of Japanese monkeys

The changes of dominance rank among female Japanese monkeys of the Koshima group over a period of 29 years from 1957 were studied. The dominance rank order was relatively stable in the early population growing phase, while large scale-changes of dominance rank order occurred successively in the phase of population decrease brought about by the severe control of artificial feeding after 1972. Nevertheless, the rank order of several females of the highest status was stable. Furthermore, the reproductive success of these highest status females was high (Mori, 1979a;Watanabe et al., in prep.). Divergence of the dominance rank order fromKawamura's rules (Kawamura, 1958) was observed in the following respects: (1) Some females significantly elevated their rank depending on the leader males. (2) If mothers died when their daughters were still juveniles or nulliparous, the dominance rank of some of these offspring females was significantly lower than the mother's one. However 55% of daughters which lost their mothers at a young age inherited the mother's rank. (3) Dominance among sisters whose mother had died when at least one of the daughters was under 6 years old followed the rule of youngest ascendancy in 60% (Kawamura, 1958), and in 80% when both of the daughters were nulliparous at the mother's death. The mean rate of aggressive interactions for each female with subordinates to her was calculated by dividing the total aggressive interactions between the female in question and her subordinates by the number of subordinate females to the female in question. A female which showed a high rate of aggressive interactions with her subordinates was categorized as an “Attacker”, and a female showing a lower rate was categorized as a “Non-attacker”. Similarly, categories of “Attacked”, and “Non-attacked” were distinguished by using the rate of aggressive interactions with dominant females. Several females which were once categorized in one category in a year were repeatedly categorized in the same category over different years. The “Attacked” tended to be females of higher rank, and “Non-attackers” tended to be females of lower rank. “The second-higher-status females”, were “Attacked”, and their rank was unstable. In particular, females of lower rank within the lineage of the highest rank suffered this kind of severe status. Most of the daughters of these females showed a sharp drop of rank, and died when they were still at a young age, i.e. “the second-higher-status females” displayed low fitness. “Non-attackers” were significantly “Non-attacked”; i.e. they were females which showed a non-social attitude. Females which underwent a drop of rank tended to be “Non-attackers”. The most important factor which determined the females' rank was the memory of their dominance relations under the influence of their mother [dependent rank (Kawai, 1958)] in their early life during development. This finding corresponds well with the results in baboons obtained byWalter (1980); the target females of aggressive interactions by adolescent females were determined by the rank of the mothers when these adolescent females were born.     

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.     

The study of vocal communication of wild mandrills in Cameroon in relation to their social structure

The vocal repertoire of the mandrill (Mandrillus sphinx), a forest living baboon, is described, and their vocal communication analyzed quantitatively. Although the vocal repertoire of mandrills corresponds well to that of savanna living baboons,Papio, some characteristics differed, such as the development of long-distance calls and differentiation of vocalizations between age-sex classes. Vocal communication within a group was closely related to changes in the spatial distribution of group members, and the two most common vocalizations, crowing and 2PG, appear to function as contact calls. Based on the wide dispersion of food trees, a group of mandrills divided into several feeding groups (subgroups). The two types of contact call were given in different and in some senses complementary contexts, and helped to facilitate and maintain group integration. According to their acoustic structure, these calls are long distance calls. Influenced by the high-level of attenuation of vocalization on the forest floor, the mandrill has developed them as contact calls, instead of using the contact “grunt,” which is common to the savanna living baboons. Comparing the patterns of vocal exchanges of mandrills with those of gelada and hamadryas baboons which have a multi-levelled society, the social structure of the mandrill is discussed. From the analysis of the spatial distribution of vocal emission, a number of clusters of vocalizations were obtained. These clusters correspond to subgroups. The frequent female-female and female-male vocal exchange between subgroups of mandrills suggest that the relationships between subgroups are less closed than between the one-male units of gelada and hamadryas baboons. Furthermore some of these clusters include more than two vocalizing adult males, while in other clusters there are no vocalizing adult males. Thus, the social structure of mandrills is suggested to be multi-male rather than a multilevelled type. The absence of contact calls specific for short distance and the functional replacement of the grunting of all group members by persistent emission of a loud call (2PG) by usually just one adult male suggests that the social structure of mandrills is not exactly equivalent to that of the multimale troop of savanna living baboons. Usually the use of 2PG is monopolized by one adult male travelling in the rear part of the group. Such monopolization of 2PG emission and the pattern of 2PG-2PG or 2PG-roar exchanges by adult males in some cases indicate the existence of strong dominance relationships among adult males, and especially the existence of a leader male within a multi-male group of mandrills.     

Ecology and sociality in a multilevel society: ecological determinants of spatial cohesion in hamadryas baboons

The multilevel society of hamadryas baboons, consisting of troops, bands, clans, and one-male units (OMUs), is commonly perceived to be an effective means of adapting to variable food availability while allowing spatial cohesion in response to predator pressure. The relationship between these variables, however, has never been tested quantitatively. The Filoha site in Awash National Park, Ethiopia is ideally suited to such an investigation as it contains nutrient-dense palm forests in addition to the Acacia scrublands typical of hamadryas distribution elsewhere, allowing comparisons of spatial cohesion across habitat types. Here, we use observations over a 1-year period to examine the relationship between resource availability, perceived predator pressure, and spatial cohesion in a band of wild hamadryas baboons at Filoha. Our results demonstrate that the band was more likely to break into OMUs when foraging in habitats with lower food availability, and that the band fissioned into independent clans more often when preferred resources were not available. Furthermore, the baboons remained in larger aggregations for longer periods of time (i.e., prior to embarking on their daily foraging route) on mornings after predators were heard in the vicinity, and increased cohesion in response to encounters with people who may have been perceived as predators. These results support the notion that hamadryas baboons change their social groupings in response to both food availability and predation risk and that the ability of hamadryas bands to cleave and coalesce in response to changes in these factors underlies the evolution of the hamadryas modular social structure.     

Social organization of Sichuan snub-nosed monkeys (Rhinopithecus roxellana) in the Qinling Mountains, Central China

Sichuan snub-nosed monkeys were observed for 197 days from 2000 to 2003 in the Qinling Mountains, Central China. The study group was provisioned in 2001 allowing detailed observations of social organization based on individual identification. The group was composed of 45–82 monkeys, all of which belonged to one of 6–8 one-male units (OMU) that foraged to form one big group. The average unit size was 9.0±2.3, 8±1.5 in the winter and 11.1±2.0 in the spring. Immigration or emigration of one-male units to or from the foraging group was observed, as was migration of individuals in and out of OMU, especially for by subadult females and juveniles. Group size therefore tended to fluctuate with the number of OMU and the number of young monkeys present in the group. The OMU in the study area were smaller than those in Shennongjia area. The factors influencing the size of these OMU and the entire group are discussed.     

Peering in mature, captive bonobos (Pan paniscus)

“Peering”—close-proximity staring at the mouth of another—was observed in ten (three males and seven females) mature (at least 7 years old) bonobos (Pan paniscus) living in three social groups at the San Diego Zoo and Wild Animal Park. Instantaneous scan samples, taken at 2-min intervals, over a three-and-a-half year period, yielded 617 observations of peering (1.4 per observation hour). Food was exchanged in only 15 of these scans. Peering was most often performed by younger animals and was primarily directed toward older females (“matrons”). In a given dyad, the animal more likely to peer at the other was also more like to both peer and be peered at if they frequently groomed and infrequently displayed aggression at a given female. An adolescent male showed the highest frequency of peering when living with two older females, but dropped to adult male levels when later housed with two younger (albeit mature) females. A reversal in which animal was more likely to peer, follow, and groom occurred in one female dyad, after the birth of the younger animal's first infant. After a similar birth in the other group, no such changes were observed. We discuss how these and related findings, in conjunction with what is known of the social structure of this species, suggest that one possible function of peering in bonobos may be as a signal acknowledging female status.