Affiliative relationships and reciprocity among adult male bonnet macaques (Macaca radiata) at Arunachala Hill, India

In captivity, male bonnet macaques (Macaca radiata) frequently express "friendship" toward one another, including affiliative behavior such as huddling, grooming, coalitionary support, and sitting in close proximity. The purpose of this study was to determine whether wild adult male bonnet macaques also express "friendship" by investigating whether or not (1) adult male bonnet macaques have affiliative social relationships with other males, (2) the strength of social relationships varies among dyads, (3) there is time-matched reciprocity in allogrooming among dyads, and if so, whether the level of reciprocity occurs within a bout of grooming, a day, or over 2 months (the limit of this study), and (4) a correlation exists between the strength of social relationships and dominance ranks among adult males. Focal samples totaling 150 hr on all seven adult males in one study group were conducted to record both agonistic and affiliative interactions. Agonistic interactions were used to construct a dominance hierarchy, whereas affiliative interactions (sitting in proximity to within 1 m with and without grooming) were used to quantify the existence and strength of social bonds within dyads. Results show that adult male bonnet macaques had differentiated affiliative relationships with other males in their group. There was little reciprocity of grooming within a bout of grooming or within a day, but greater reciprocity over the study period of 2 months. There was no correlation between dominance ranking distance and the strength of affiliative relationship within dyads; however, within dyads lower-ranking males groomed higher-ranking males more than vice versa. This study suggests that friendships in male bonnet macaques are characterized not by immediate tit-for-tat reciprocal altruism, but by reciprocity over a longer time span, and that affiliative social relationships may be less constrained by agonistic relationships than is the case in more despotic species of macaques.     

Social Behavior,Reproductive Strategies,and Population Genetic Structure of <Emphasis Type="Italic">Lagothrix poeppigii</Emphasis>

For species of primates in which females emigrate, we would expect males within groups to be related to one another. Kin selection theory suggests that these males should associate preferentially with one another, be more affiliative and cooperative with one another than females are, and compete less overtly with one another over reproductive opportunities than males in female philopatric taxa do. Precisely these patterns of social behavior characterize well-studied populations of 2 of the 3 atelin primate genera: spider monkeys (Ateles) and muriquis (Brachyteles). For the third atelin genus, Lagothrix, patterns of intragroup social behavior have been less well-documented. We studied the social and reproductive behavior of lowland woolly monkeys (Lagothrix lagotricha poeppigii) in Ecuador during a one-year observational study and subsequently used molecular techniques to investigate population genetic structure and dispersal patterns for this taxon. Among adult male woolly monkeys, both affiliative and agonistic interactions were rare, and males were seldom in close proximity to one another. Relationships among male woolly monkeys are best characterized as tolerant, especially in the context of mating wherein direct competition among males was minimal despite the fact that females mated with multiple males. Relationships among females were likewise generally tolerant but nonaffiliative, though females often directed harassment towards copulating pairs. Affiliative interactions that did occur among woolly monkeys tended to be directed either between the sexes—primarily from female to male—or from younger towards older males, and the proximity partners of females tended to be members of the opposite sex. These results suggest that bonds between the sexes may be more important than same-sex social relationships and that direct female-female competition is an important feature of woolly monkey reproductive biology. Our genetic results indicate that, as in other atelins, dispersal by females is common, but some male dispersal likely occurs as well. In some but not all groups we studied, nonjuvenile males within social groups were more closely related to one another on average than females were, which is consistent with greater male than female philopatry. However, differences in these patterns among our study groups may reflect local variation in dispersal behavior.     

Male-female affiliative relationships in naturally occurring ringtailed lemurs (Lemur catta) at the Beza-Mahafaly Reserve,Madagascar

Affiliative behavior between adult male and female ringtailed lemurs was examined as part of a project concerning male affiliation with conspecifics of all age/sex classes. Males in three social groups were studied over a 12 month period. Male-female preferred partnerships existed, and were variable according to reproductive season. Dominance rank, age, or tenure of the male did not appear to affect either the number of partnerships or frequency of affiliative behaviors that males had with females. However, males residing in groups with fewer males exhibited both higher frequencies of affiliative interactions with females and were nearest neighbors to females more often than males living in a group containing more males. Females were found to be responsible for proximity maintenance of male-female dyads in the majority of cases. Neither reproductive season nor seasonal availability of food resources strongly affected the frequency of affiliative interactions between males and females, it is proposed that an important aspect of successful group membership for male ringtailed lemurs relates to the development of social relationships with adult females. Males can benefit from such relationships in terms of greater centrality to the spatial core of the group, which can result in enhanced predator protection, greater opportunities for social contact, and potentially greater access to estrous females. © 1996 Wiley-Liss, Inc.     

Effect of Male Emigration on the Vigilance Behavior of Coresident Males in White-Faced Capuchins (Cebus capucinus)

I examined the vigilance behavior of male Cebus capucinus residing in four groups in Santa Rosa Park, Costa Rica. One male emigrated from each of three study groups, providing ideal experimental conditions for examining the effect of coresident males on male vigilance behavior (social and non-social). Following the predator and conspecific defense theories for the adaptive value of male vigilance behavior and the occurrence of multimale groups, I predicted that male nonsocial vigilance would increase after the emigration of a coresident male. My prediction was supported in only one of the three study groups. Males in two groups decreased their nonsocial vigilance after the emigration of a coresident male, which was probably influenced by seasonal changes and the peripherality of one group's coresident male before his emigration. The social vigilance hypothesis proposes that for species with rigid dominance hierarchies, social vigilance should increase with an increase in same sex competitors. Therefore, I predicted that male social vigilance would decrease after the emigration of a coresident male. The hypothesis is generally supported in my findings, as two of the three groups displayed a decrease in social vigilance after the emigration of a coresident male. Increased social vigilance in the third group is best explained by the change in the alpha male's social behavior after the emigration of his only coresident male.     

Male chimpanzee behavior in relation to female ano-genital swelling

This study examined the relationships between male agonistic, affiliative, and sexual behaviors and female estrus condition in captive adolescent and young-adult chimpanzees (Pan troglodytes). Data on agonistic, affiliative, and sexual behaviors of 11 males living in three social groups were collected during daily 45 minute observations over a 5 month period. Female estrus condition was assessed daily using the relative size of the female's ano-genital swelling. It was hypothesized that the presence of maximally tumescent females would generate conflicts between males, so an increase in inter-male agonism was predicted. Males exhibited higher rates of agonism toward other males when at least one female in the group was maximally tumescent. Male affiliative behavior directed toward other males and social play with males were affected by the presence and number of maximally tumescent females. Male sexual behavior increased when maximally tumescent females were present.     

Age related variation in male–male relationships in wild spider monkeys (Ateles geoffroyi yucatanensis)

In social organizations characterized by male philopatry, social relationships between males are argued to be the strongest. Little is known about the social relationships of philopatric male spider monkeys. To address this limitation, we investigated social relationships among individually recognized wild adult male spider monkeys from two well-habituated communities in the Yucatan Peninsula, Mexico, focusing on affiliative behaviors important in regulating male social relationships, including grooming, embracing, arm-wrapping, and grappling. We examined whether behaviors were reciprocated between male partners and whether age was a factor in how the behaviors were distributed or reciprocated, by examining differences between younger adult males (<10 years) and older adult males (≥14 years). Although we found evidence that affiliative behaviors were overall reciprocated between spider monkey adult males, there were pronounced differences in the interactions depending on their relative age. Reciprocation in grooming and embraces between same-age males suggests their relationships are valuable to both partners. Among different-age dyads, younger males gave more embraces than they received, were the initiators of grappling and arm-wrapped more often than with same-age males, suggesting relationships between younger and older males are more risky. This confirms that younger males are attracted to older males, probably because they value relationships with older males more than the reverse, but they are also at risk.     

Social behavior in fission–fusion groups of red uakari monkeys (Cacajao calvus ucayalii)

Primates living in large groups that divide to forage must have social systems compatible with this mode of living. Uakari monkeys (Cacajao spp.) live in large groups and exhibit a form of fission–fusion grouping, but their social organization is poorly understood. We present some of the first data on social behavior for this genus based on a study on Cacajao calvus ucayalii. They traveled in multimale multifemale groups of highly variable sizes, with bachelor units on the periphery. Adult males were affiliative, and adult females associated with more than one adult male. Adult females typically traveled with their dependent offspring and an older juvenile within the group. In parties of two or more males, individuals engaged in previously unreported display behaviors and acted together to aggressively chase other males. Breeding was seasonal, and mating occurred away from other group members. We speculate on the social organization of C. calvus ucayalii, in which dispersal may be bisexual and peripheral males are affiliative with one another. Affiliated males appear to cooperate in fighting and displaying to other males for access to females during the breeding season. Am. J. Primatol. 71:976–987, 2009. © 2009 Wiley‐Liss, Inc.     

Reconciliation in Male Stump‐tailed Macaques (Macaca arctoides): Intolerant Males Care for Their Social Relationships

Recent research has shown that social relationships may exert positive effects on fitness. Therefore, it is expected that animals make efforts to develop and preserve close social bonds. Reconciliation is a conflict resolution mechanism, which allows buffering the negative effects of aggressive conflict on social relationships. As socioecological theories claim that intrasexual competition and conflict‐shaped social interactions among macaque males, postconflict behaviour should reflect the effects of these evolutionary forces. Here, we present the results of a study on a provisioned group of Macaca arctoides in Thailand focusing exclusively on male postconflict behaviour. Through an Information Theory–Model Selection approach, we evaluated the relative support for three predictions about the occurrence of reconciliation and the quality of the opponents' social relationships. The strength of the opponents' affiliative relationship was the only variable predicting reconciliation occurrence. This agrees with the main prediction of the valuable relationship hypothesis for the function and distribution of reconciliation, which states that the opponents sharing high‐quality relationships are more likely to reconcile, and this interaction is aimed to repair the disturbing effects of previous aggression on their relationship. The conciliatory tendency found among these wild males is much lower than reported previously for this species from captive conditions, matching more closely the rates reported for species with an intolerant‐despotic dominance style. The results offer strong support for the importance of developing and preserving valuable relationships, even among rather intolerant males.     

Bridging behavior and other affiliative interactions among male tibetan macaques (Macaca thibetana)

I describe bridging behavior and social relationships between adult males and infants in a free- ranging group of Tibetan macaques (Macaca thibetana)at Mt. Huangshan, China. The subjects performed bridging in which two adult males simultaneously lifted up an infant, sucked or touched its genitalia, and then groomed each other in nonagonistic contexts. Males also expressed social behaviors with other males, such as mounting, penis-sucking, and embracing while touching each other’s penes. Males also employ bridging while exploiting an infant as a social tool, not only to reduce the probability of an aggressive response from dominant males (agonistic buffering), but also to develop and to maintain affiliative social relationships with other males. Use of male infants in bridging contributed to frequent male-infant interactions such as holding,grooming, and penis-sucking. Although these interactions might not have a positive influence on infant survival, they may facilitate the maintenance of affiliative relationships with adult males until they reach maturity. The development of bridging might have a close relation to the high socionomic sex ratio (adult male/adult female) and frequent affiliative interactions between males, especially among the adolescents and adults.     

Sex differences in the social behavior of wild spider monkeys (Ateles geoffroyi yucatanensis)

Competition for resources and the need for cooperation are reported to affect patterns of social interactions and thus the quality of social relationships in primates. Relationships may be described as high quality when both individuals behave in a way that benefits their partner. We investigated the distribution of a wide range of social behaviors across sex partner combinations of adult spider monkeys with specific reference to contested resources. Data were collected from two communities of wild spider monkeys (Ateles geoffroyi yucatanensis) in the Otoch Ma'ax Yetel Kooh reserve in Yucatan, Mexico. Affiliative behavior was exchanged most frequently between males, and as male-male aggression was rare, male-male social relationships were characterized as high quality. Female-female social relationships were best described as low quality as females showed no preference to be in proximity with other females and female-female affiliative behavior was rare. Relationships between the sexes generally were characterized by high rates of female-directed male aggression, although additional investigation into the effects of kinship and female reproductive state on male-female relationships is required before further conclusions can be drawn. Dyadic and coalitionary female-female aggression was significantly higher than expected in a feeding context, confirming that female spider monkeys primarily compete for access to food. Male-male aggression did not increase significantly when in the presence of females, but males embraced at higher rates when in mixed-sex subgroups. As embraces serve to reduce the likelihood of aggression during tense situations, high rates of male-male embraces in mixed-sex subgroups may mitigate male conflicts over access to females.     

Social compatibility in a newly formed all‐male group of white crowned mangabeys (Cercocebus atys lunulatus)

Surplus males in primate captive populations are a common problem for zoos. Some captive breeding programs promote all‐male groups as an adequate option to house surplus males, but there have been few attempts to assess the feasibility of this management technique across primate species. The present study provides preliminary data regarding social compatibility within a newly formed all‐male group of four white crowned mangabeys (Cercocebus atys lunulatus). The study was conducted at the Valencia Zoo (Spain), where data on social behavior were collected over 6 months using continuous focal animal sampling for a total of 87 hr of observation. Results show that low intensity aggressive behaviors (facial threats) were expressed at high rates, whereas physical aggression (fights) rarely occurred. Aggression was more frequent among individuals belonging to the same age–gender class. Regarding affiliative behaviors, every individual actively sought proximity to all other group members through positive approaches, and although not all males carried out social grooming, every male was groomed by at least one group member. Our results suggest that the group was compatible socially because social relationships among the individuals were not neutral, and physical aggression occurred at low rates. The present study provides preliminary data supporting the feasibility of all‐male groups as a management option for surplus males in captive populations of white crowned mangabeys. Nevertheless, further studies are needed to be able to generalize both within and across species. Zoo Biol 0:1–7, 2007. © 2006 Wiley‐Liss, Inc.     

Intragroup Social Relationships of Male Alouatta palliata on Barro Colorado Island,Republic of Panama

We examined social and spatial relations of adult males in one group of mantled howling monkeys (Alouatta palliata) on Barro Colorado Island (BCI) in central Panama to document patterns of association. Beyond the existence of an alpha male, we could not distinguish any linear dominance hierarchy among the 6 study males. All males copulated with estrous females. Our findings contrast with reports of intragroup male behavior in Costa Rican howlers. Study males engaged in little or no affiliative or agonistic behavior with one another, but engaged in significantly more such interactions with females. The alpha male, the oldest male and a younger male were most frequently in association with females. Of group males, the oldest male associated significantly more with other males. Overall, male behavior in mantled howling monkeys on BCI generally followed the van Hooff and van Schaik (1994) model of male relationships. The low incidence of intragroup social interactions of any type in the focal males may reflect the energetic costs of social behavior. We suggest that intragroup social relationships among mantled howler males are structured by more subtle means than overt physical interactions, possibly including vocal communication, relationships with individual group females, and kinship.     

Social boundaries in a Malagasy Prosimian,the Sifaka (Propithecus verreauxi)

Using focal animal samples, the social organization of sifakas was studied in two forests for 2500 hr spread over 18 months. Data were also obtained on the size and composition of groups at two other sites. The size and adult sex ratio of groups varied widely within populations, although population-wide sex ratios approached unity. During the brief annual mating season, some males mated with females belonging to other groups. The response of both male and female group members to the approach of a nongroup male appeared to depend on his identity. It is argued that while group size may be constrained by the availability of resources, group composition reflects the effects of small-sample demographic variations rather than the outcome of reproductive competition among or between males and females. Spatial boundaries of groups do not coincide with social or reproductive boundaries even within a single breeding season; further study is needed of the processes whereby relationships are established and maintained between individuals from different groups.     

Male affiliation, cooperation and kinship in wild chimpanzees

Long-term field research has revealed that male chimpanzees, Pan troglodytes, affiliate and cooperate in several contexts. Assuming close genetic relationship among males, affiliative and cooperative behaviour have been hypothesized to evolve through the indirect effects of kin selection. We tested the hypothesis that matrilineal genetic relatedness affects patterns of male social affiliation and cooperation in an unusually large community of chimpanzees at the Ngogo study site, Kibale National Park, Uganda. Field observations indicated that six behavioural measures of affiliation and cooperation among 23 adult males were significantly correlated with each other. Sequences of the first hypervariable portion of the mtDNA genome revealed that three pairs of males and one quintet shared mtDNA haplotypes. Matrix permutation tests using behavioural and genetic data showed that males that affiliated and cooperated with each other were not closely related through the maternal line. These findings add to a growing body of empirical evidence that suggest kinship plays an ancillary role in structuring patterns of wild chimpanzee behaviour within social groups. Copyright 2000 The Association for the Study of Animal Behaviour.     

Integration of New Males into Four Social Groups of Tufted Capuchins (Cebus apella)

We examined how aggressive, affiliative, and sexual behavior function to integrate male capuchins (Cebus apella) into a new social group. Nine males were exchanged among four social groups. We performed instantaneous scans and all-occurrence sampling during baseline, introduction, and follow-up periods. The study included three different introduction situations: 1) males familiar to one another were introduced to a group with no other adult male, 2) males unfamiliar to one another were introduced to a group with no other adult male, and 3) males familiar to one another were introduced to a group with an existing elderly, resident male. Severe aggression occurred in situations 2 and 3, but the introductions were peaceful in situation 1. In all cases proceptive females were among the first individuals to affiliate with the males, and males did not appear to compete for access to proceptive females. Following their period of proceptivity, the females that had cycled remained preferred social partners for the males. Immature animals also quickly affiliated with the new males, and the males tolerated the attention from immatures. Affiliative relationships between the males and nonproceptive females developed slowly, and while male-female aggression was mild, aggression among adult males (familiar and unfamiliar) had the potential to be severe.     

Within-group social relationships among females and adult males in wild western lowland gorillas (Gorilla gorilla gorilla)

Within-group female relationships, and relationships between females and adult males have not been described for wild western lowland gorillas. New data are presented here from Mbeli Bai in the Nouabale-Ndoki National Park, Republic of Congo. Nine groups were observed over a continuous 2-year period, and affiliative and agonistic interactions, as well as time spent by females and adult males in proximity to each other were recorded. Affiliative behavior was extremely rare in the bai, and measures of proximity between individuals were inconclusive because of the small sample sizes. Rates of agonistic interaction among females were highly variable between groups. The majority of agonistic interactions were undecided, and the supplant rate was low. Evidence for an agonistic dominance hierarchy is weak. The data do not support hypotheses that increased frugivory in western lowland gorillas raises levels of within-group contest competition; however, the need for intersite comparisons is stressed. In contrast, rates of agonistic behavior between females and silverback males were higher and more consistent between groups. Agonistic relationships between males and females conformed with patterns seen in mountain gorillas, where male protection from intraspecific aggression is important. The benefit females gain by forming long-term cooperative alliances appears to be small, and is likely outweighed by reproductive advantages associated with male protection.     

Reunion displays in captive male brown capuchins (Cebus apella)

Separation and subsequent reintroduction of brown capuchins (Cebus apella) can lead to reunion displays during which individuals will embrace and emit high-pitched, loud vocalizations. These displays were investigated in three male capuchins: two adults and one juvenile. These subjects were part of a larger captive social group held at the Capuchin Laboratory, Hiram College, OH, USA. We hypothesized that males would engage in reunion displays following separation regardless of the order of reintroduction, as evidence of an affiliative bond judged by concurrent social behavior. Sixty separations were conducted as part of weekly husbandry over a period of 13 months. Occurrence of reunion displays between the males was recorded with the initiator also noted. Reunion displays among males were found to occur only between the juvenile and alpha male, and all displays were initiated by the juvenile. Displays were highly influenced by the order of reintroduction. We suggest that reunion displays operate to reaffirm social relationships, particularly during situations of social tension.     

Social interactions among captive brushtail possums (Trichosurus vulpecula)

A biological control strategy relying on a self-disseminating agent may provide the only affordable long-term technique for reducing brushtail possum (Trichosurus vulpecula) numbers throughout New Zealand. The objective of this study was to determine the frequency and patterns of social interactions in single and mixed-sex groups of possums, in order to identify interactions that may assist in the dissemination of biocontrol agents. Thirty-two sexually mature wild-caught possums (16 female, 16 male) were housed in captive groups (of four) in enclosures during the breeding and non-breeding seasons. Groups consisted of either all females, females and males or all males. Four types of social interactions were defined: threats, fights, affiliative and sexual interactions. Interactions only occurred between two animals at one time and were most frequent in the non-breeding season. Within all female groups, possums readily engaged in interactions that had either an agonistic or affiliative nature. Mixed-sex groups interacted less frequently than females, but also engaged in both agonistic and affiliative interactions. In contrast, possums in all male groups rarely interacted, with only a few fights and no affiliative interactions observed. Some mixed-sex dyads appeared to 'associate' during the breeding period. Young were produced by three females that regularly associated with a male and one female that showed little associative behaviour. Given the different interaction patterns observed in each group type, biological control agents that rely on specific interaction patterns for dissemination, are likely to spread at different rates among different possum groups and in different seasons.     

Social and hormonal mechanisms underlying male reproductive strategies in black howler monkeys (Alouatta pigra)

We investigated the social and hormonal mechanisms underlying male reproductive strategies in two multimale-multifemale groups of black howler monkeys (Alouatta pigra) during a 14-month study in Palenque National Park, Mexico. Fecal glucocorticoid (fGC) and androgen (fA) levels were analyzed for 343 fecal samples collected from 14 males during their presence in the study groups. Neither immigrating males nor resident males that remained in the group had elevated fGC and fA levels during 11 observed male migration events, suggesting that competition over group membership was not associated with variation in the fecal hormonal levels of males. Instead, fGC and fA levels were significantly higher in males who maintained a central position in the group and had almost exclusive access to fertile females than in other resident males. These “central” males were responsible for maintaining close spatial associations and cultivating strong affiliative relationships with cycling, sexually active females but not with noncycling, sexually inactive females. “Noncentral” males did not form strong social relationships with either cycling or noncycling females and had no or very few mating opportunities. Our findings suggest that male black howler monkeys compete nonaggressively by fostering relationships with cycling females and that the elevated fGC levels of central males may be indicative of the social challenges involved in their indirect competition.     

Male-male Interactions in Heterosexual and All-male Wild Mountain Gorilla Groups

Variation in male dispersal and behavior patterns are components of intraspecific differences in social systems. A comparison of male behavior in different social settings can be useful for determining which behavioral mechanisms contribute to variability in social systems. Two heterosexual multimale groups and one all-male group of mountain gorillas (Gorilla gorilla beringei) were observed for over 1100 h at the Karisoke Research Centre, Rwanda. Data on proximity patterns, dominance relationships, aggression, agonistic interventions, affiliation, and homosexual behavior were compared among the males in these groups to examine the influence of female presence, sex ratio, group size, and kinship on male—male interactions. Males in the all-male group stayed closer together, affiliated more, exhibited more homosexual behavior, and were more aggressive toward each other than males in heterosexual groups. However, the males in heterosexual groups showed more wounding and more consistent dominance relationships. Kinship did not influence male-male relationships in the all-male group. The males in the heterosexual groups rarely interacted with one another; they may actively avoid close proximity to reduce aggression. Results suggest that the variable social system of mountain gorillas may be more strongly influenced by demographic factors, male-female social relationships, and male-male competition for mates than by any benefits of male-male relationships.