Agonistic Interactions and Matrifocal Dominance Rank of Wild Bonobos (Pan paniscus) at Wamba

I studied dominance relations in a wild group of bonobos at Wamba, Democratic Republic of Congo. Although agonistic interactions between males occurred frequently, most of them consisted only of display, and physical attacks were infrequent. Dominance rank order seemed to exist among males, but its linearity is unclear. Dominant males rarely disturbed copulatory behavior by subordinate males. However, high-ranking males usually stayed in the central position of the mixed party and, so, would have more chance of access to estrous females. Among females, older individuals tended to be dominant over younger individuals. However, agonistic interactions between females occurred rather infrequently, and most consisted of displacement without any overt aggressive behavior. Dominance between males and females is unclear, but females tended to have priority of access to food. The close social status between males and females may be related to the prolonged estrus of females and their close aggregation during ranging. Existence of a male's mother in the group and her dominance status among females seemed to influence his dominance rank among males. Young adult males whose mothers were alive in the group tended to have high status. In some cases, change in dominance between high-ranking males was preceded by a corresponding change in dominance between their mothers. As the dominance status of females is similar to that of males, mothers may be able to support their sons to achieve high status, stay in the center of the mixed party, and so have greater access to females, which may maximize the number of descendants of the mothers.     

Party composition and dynamics inPan paniscus

The pygmy chimpanzee, or bonobo, Pan paniscus,diplays a fission-fusion social organization in which individuals associate in parties that vary in size and composition. Data from a 2-year field study of nonprovisioned P. paniscusshow that party composition varies with party size. Although females, on average, outnumber males, the proportion of males in the party increases in larger parties. This effect was not due to the greater number of known females. Both females and males will join and leave a party in the company of others, but only males appear frequently to join or leave as lone individuals. All-male parties were not observed, but all-female (nonnursery) parties were relatively common. These trends reflect greater cohesion among females than observed in P. troglodytes schweinfurthii.Cohesion between males and female P. paniscusmay increase with party size.     

Sex and group differences in feeding on animals by wild chimpanzees in the Mahale Mountains National Park,Tanzania

Sex differences in animal prey intake were revealed by fecal analysis among wild chimpanzees of the large-sized M-group (ca. 100 members) in the Mahale Mountains National Park, Tanzania: prime adult or old males feed more on vertebrates, while adult females more onCamponotus ants. By contrast, such differences were not obvious in the neighboring, small-sized K-group (ca. 20–30 members), despite the similar environment in which the two unit-groups lived. Such sex and group differences may be explained in terms of various factors, either ecological or social, or both, but social factors seem most responsible in particular for the group differences. It seems likely that increased capture rate of vertebrates per unit-group in the larger-sized M-group results in increased per capita intake of meat among prime adult or old males. Also, the more frequent interactions among prime adult or old males of M-group appear to reduce the frequency of theirCamponotus ant-fishing behavior.     

Association,ranging, and the social unit in chimpanzees of the Mahale Mountains,Tanzania

In order to characterize the social unit in chimpanzees, about which several conflicting views have been proposed, the proximity matrix among 55 recognized chimpanzees and the range covered by each of them are examined, on the basis of data obtained at the Mahale Mountains during 12 months in 1978–1979. It is shown once again that chimpanzees have a bisexual social unit (unit-group). Two such unit-groups were detected in the study area. All animals belonged to one of the two unit-groups except a few cycling females (and a juvenile male accompanied by his cycling mother) which were seen to associate alternately with members of two neighboring unit-groups, covering a whole range of one or even two unit-groups. The problem of such females is discussed in relation to the spatial relationships between the two unit-groups. Reexamining the membership of a unit-group, it is demonstrated that a unit-group was most likely patrilineal. While nulliparous females transferred between unit-groups, parous females tended to remain in a unit-group where they first gave birth to infants and to have several offspring therein. This appeared significant for ensuring recruitment of members of the next generation to a patrilineal unit-group. Although some adult males left their natal unit-group, they never joined the other. Male departure from a unit-group seemed to be forced by the other males and to be the sociological equivalent of going into exile, which is unique in nonhuman primates.     

Birth-season interactions of adult female Japanese Macaques(Macaca fuscata) without newborn infants

The affiliative interactions of 11 adult female Japanese macaques that did not deliver an infant during the 1981 birth season of the Arashiyama West troop were examined. Consideration was given to the effects of kinship as a structuring element in these birth-season interactions and to the degree of association with various categories of troop members based on age, sex, and (in the case of adult females) whether or not the females were new mothers. Females without infants interacted predominantly with their yearling off-spring, although it was the behavior of the offspring that precipitated the interaction. These females were active in soliciting affiliation with nonkin new mothers, whereas female matrilineal relatives with new infants approached and remained in proximity to them more than did nonrelated new mothers. Females without newborns groomed and approached nonkin infants more than infants within their own matriline, and these infants were predominantly those of females in the highest-ranking matriline of the troop. Adult males were responsible for 40% of all grooming received from nonkin by the females without newborns, and these males approached them significantly more than did other adult females without infants. These patterns demonstrate that the structure of social relationships is influenced by the particular dynamics of troop contexts such as birth seasons, as well as by enduring, broad-based affinities which are less affected by cyclic changes in troop context.     

Male-male relationships among wild bonobos (Pan paniscus) at Wamba,Republic of Zaire

Male-male relationships among wild bonobos (Pan paniscus) in two adjacent unitgroups (E1 and E2 groups), which were formed by division of the E group, were studied at Wamba, in the Central Zaire Basin, by analyzing the proximity and social interactions among males. Dominant-subordinate relationships between a male-male dyad were easily recognized from the directions of individual agonistic interactions. Male bonobos rarely joined forces in aggression. Clear differences in social status existed between adult and adolescent male bonobos in both groups, as reported in the case of chimpanzees (Pan troglodytes). The presence of mothers in the unit-group greatly influenced the dominant-subordinate relationships among males through strong mother-son bonds in both groups. However, the extent of the mother-son bonds differed between the groups. Males in the E2 group participated more frequently in agonistic or affinitive interactions than did males in the E1 group. Males in the E1 group were divided spatially into several clusters, while there were cohesive relationships among the adult males in the E2 group. The difference in intensities of mother-son bonds between the groups may be explained by the distribution of males at the time of the division of the E group. Differences in male-male relationships between bonobos and chimpanzees seem to be related to differences in intra- and inter-unit-group competition among males between the two species. Male chimpanzees may achieve coexistence by manipulating ambivalent relationships that are caused by intra- and inter-unit-group competition among them, while male bonobos may achieve coexistence by decreasing intra- and inter-unit-group competition among them.     

Seasonal variation in association patterns of wild spider monkeys (<Emphasis Type="Italic">Ateles belzebuth belzebuth</Emphasis>) at La Macarena,Colombia

Spider monkeys exhibit a fission–fusion type of social organization. I studied party size and party composition in wild long-haired spider monkeys (Ateles belzebuth belzebuth) in three study periods at La Macarena, Colombia and found that overall party size was larger in the fruit-abundant season. Mean party size in which males were observed was relatively stable across seasons. In contrast, the mean party size of females varied. Females were observed in larger parties in the fruit-abundant season than in the fruit-scarce season. Moreover, whereas males associated with each other at an almost equal frequency across seasons, females associated with each other more frequently in the fruit-abundant season. Females with infants or small juveniles were more often in association with other individuals than were cycling females. The intensity of individual relationships varied according to season, such that even mothers and sons were not always strongly associated. In a large party, females with infants may gain from predation avoidance but they are at a disadvantage in terms of scramble competition. The balance between these factors may change with fruit availability and may influence party size in different periods. For males, party formation may facilitate the defense of resources from neighboring groups more than provide predation avoidance. Electronic Publication     

Group unity of chimpanzees elucidated by comparison of sex differences in short-range interactions in Mahale Mountains National Park,Tanzania

Chimpanzees (Pan troglodytes) form multi-male and multi-female unit groups with fission–fusion grouping patterns. Short-range interaction (SRI) plays an important role in the unity of these groups and in maintaining social bonds among members. This study evaluated three models of chimpanzee social structure that differed according to the emphasis each placed on social bonds between the sexes, i.e., the male-only, the bisexual, and the male-bonded unit-group model. I investigated differences in SRI between the sexes among group members in well-habituated wild chimpanzees in Mahale Mountains National Park, Tanzania. I followed six focal adult males and six females, and quantified their respective SRI with other chimpanzees. Except between subordinate males and adult females, adults in general engaged in SRI with about 60–90% of the individuals with whom they made visual contact each day, whether in large or small parties. Although the number of social grooming (SGR) partners was limited, male–male SGR networks were wider than were either male–female or female–female SGR networks among adults. The number of contact-seeking behavior (CSB) partners was also limited, but dominant males had more CSB partners. Adult females mainly interacted by pant-grunt greeting (PGG) with adult males, but tended to do so mainly with the highest-ranking male(s) within visual contact. These results indicated that the social bonds among adult males were essential to group unity. Because of clear male dominance, adult females established peaceful coexistence with all group members despite less frequent SRI with subordinate males by maintaining affiliative social bonds with dominant males, thereby supporting the male-bonded unit-group model. Adult females had many female SRI partners, but these interactions did not involve performing conspicuous behaviors, suggesting that females maintain social bonds with other females in ways that differ from how such bonds are maintained with and between adult males.     

Social structures in Pan paniscus: testing the female bonding hypothesis

Based on previous research in captivity, bonobos, Pan paniscus, have been called a female-bonded species. However, genetic and behavioural data indicate that wild females migrate. Bonding between these unrelated females would then be in contradiction with socio-ecological models. It has been argued that female bonding has been overemphasized in captive bonobos. We examine patterns of proximity, grooming and support behaviour in six well established captive groups of bonobos. We find that female bonding was not a typical characteristic of all captive bonobo groups. In only two groups there was a trend for females to prefer proximity with other females over association with males. We found no evidence that following or grooming between females was more frequent than between males and unrelated females or between males. Only in coalitions, females supported each other more than male–female or male–male dyads. We also investigated five mother–son pairs. Grooming was more frequent among mothers and sons than in any other dyad, but sons did not groom their mothers more than males groomed unrelated females. Mothers groomed their sons, or provided more support to them than females groomed or supported unrelated males. Thus, while bonds between females were clearly present, intersexual relations between males and either unrelated females or their mothers are of more, or equal importance.     

A preliminary study of selective visual attention in female mountain gorillas (Gorilla gorilla beringei)

Visually attending to conspecifics can give group-living primates important ecological information, help them to anticipate the behavior of others and to regulate interactions with them, and provide other valuable social information. Variation in the importance and quality of social relationships should influence the way individuals selectively attend to fellow group members. Preliminary data on visual monitoring of conspecifics by wild female mountain gorillas (Gorilla gorilla beringei) show that selective attention mirrors variation in social relationships. Social bonds between males and females are central to gorilla society; correspondingly, females are more likely to stop feeding, and focus their attention on males who walk into view than on females, especially when males give dislays. Females are more likely to focus on other females with whom they have antagonistic relationships than those (mostly close relatives) with whom they have affiliative, cooperative ones. Further research on the context and consequences of visual monitoring could help to address questions about the regulation of social relationships and about social cognition in gorillas.     

Chemosensory discrimination of male age by female Psammodromus algirus lizards based on femoral secretions and feces

Chemical communication plays an essential role in several social and reproductive behaviors of many animals. In lizards, the main sources of semiochemicals are femoral or pre‐anal gland secretions and feces. In male lizards Psammodromus algirus, there are age‐related differences in the chemical composition of femoral gland secretions and in the reproductive strategies, with older males defending territories and females, while younger males adopting sneak‐mating strategies. Females flee more often from mating advances of young males than from those of old males, which are more successful in obtaining matings. This suggests that age discrimination of males may be important for females. We tested here whether females showed differential chemosensory responses to chemical cues (femoral gland secretion and feces) of males of two age classes, and whether females use information from substrate scent marks of males of different ages to select where to stay. We found that females elicited more tongue‐flicks to the secretion and feces of old males than to control or secretion and feces of young males. Also, the time spent by females on a scented paper depended on the treatment, suggesting that females tended to spend more time on scent marks made with femoral secretions of old males. Adult females seemed capable to discriminate between young and old males based on chemical cues alone and showed more interest in scents of old males. However, substrate scent marks did not seem to entirely determine site selection by females, suggesting that females might need additional cues to perform the choice. These results can be explained by the different age‐dependent reproductive strategies of males, which can affect differentially to females.     

Spatial structure of chacma baboon groups

A method is presented for measuring the location of individuals with respect to the center or perimeter of a primate group. The method estimates an individual’s domain of danger: the area in which a hidden predator is closer to the individual than to other group members (cf. Hamilton, 1971). A domain is determined by the directions and distances of particular neighbors from a given individual. Animals at peripheral locations have relatively large domains, whereas animals at central locations have relatively small domains. Domains of danger were sampled for members of two groups of chacma baboons (Papio ursinus)in northern Botswana, to test for nonrandom spatial patterns throughout the day. Females with infants had significantly smaller domains than did females without infants. Domains also were correlated with social ranks of females;high-ranking females had smaller domains than did low-ranking females, possibly because high-ranking females were more likely to have infant offspring. For adult males, however,domain sizes were not significantly correlated with social ranks. Immigration status of adult males, rather than social rank, better accounted for spatial positioning.     

Age differences in social interactions of young males in a chimpanzee unit-group at the Mahale Mountains National Park,Tanzania

The behavior and social interactions of young male chimpanzees were studied in relation to their age change. The data were obtained at the Mahale Mountains National Park, during a four-month period in 1986. Early adolescent males, becoming independent of their mothers, spend a long time near adults of both sexes. Late adolescent males are not tolerated by the senior males. Although such animals do not stop traveling together with their seniors, they are separated from the other members including the males of their own age class, and each of them lives a relatively lonely life. Where seniors are not nearby, they perform charging displays in front of estrous females. Young adult males tend to remain in the proximity of the alpha male, and can associate with their seniors without pant-grunting. Although some young adult males dominate over some senior males, increasingly performing charging displays, they do not appear to be permitted to associate intimately with their seniors; they are not yet considered to have attained social maturity. Prime and senior males are strongly bonded with one another, being able to associate intimately with those of their own or senior age classes including the alpha male. A young adult male's rise in rank is not connected with joining the “adult male-cluster,” nor does a senior male's decline necessarily means his dropping out from the cluster: the social position of male chimpanzees cannot be understood solely from their agonistic dominance rank. The alpha male plays a leading part in integrating the males of the unit-group. Young adult males and their seniors tend to associate most frequently with him, and all the males of the early adolescent or senior age classes pay attention to his movements.     

Proximal Spacing Between Individuals in a Group of Woolly Monkeys (Lagothrix lagotricha) in Tinigua National Park, Colombia

I studied proximal spacing within a group of woolly monkeys (Lagothrix lagotricha) during 7 months at Parque Nacional Tinigua, Colombia. I collected a total of 1188 instantaneous samples on focal individuals, recording the number and age/sex class of individuals that were in contact with, <2 m from, <5 m from the focal animal. The results indicate that proximate spacing reflects social affinities and is related to mother–infant relationship and social grooming. Subadult females and adult males are the sex/age classes with the lowest number of individuals in proximity. There are low proximity between adult females and between adult males and high frequencies of nearness between mother and offspring. Associations between males and females were usually low, but in some cases males showed preferences for a given female. There was a relatively gradual increase in spacing between mothers and their offspring as they became older. Old juvenile males were associated chiefly with other males—mostly subadults—whereas juvenile females maintained some proximity only to their mothers. There are also differences in spacing behavior according to different activity types.     

Sexual swelling,receptivity, and grouping of wild pygmy chimpanzee females at Wamba,Zaïre

The sexual swelling and copulatory behavior of ten pygmy chimpanzee (Pan paniscus) females in a wild group were studied at Wamba, Republic of Zaïre. Since the maximum size of the sexual skin revealed great individual variations depending on age and adolescent females showed little cyclic fluctuation in size, the sexual swelling was measured according to its firmness which periodically fluctuated in all age classes. The duration of maximum swelling and the cycle length were longer inP. paniscus than inP. troglodytes. Although pregnant females and those with newborn infants were sexually inactive, females with infants older than 3 years copulated as frequently as those without dependent infants. Contrary to previous reports on the sexuality ofP. paniscus, copulation was mostly restricted to the maximum swelling phase. All females were usually found in a large mixed party containing both sexes and offspring, regardless of their sexual receptivity.     

Grouping Patterns and Competition Among Female <Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis> at Ngogo,Kibale National Park,Uganda

In mammals, access to mates is probably the most important influence on male reproductive success, whereas foraging efficiency is probably the most important influence on female reproductive success Emlen and Oring (Science 197:215–223, 1977). Male chimpanzees (Pan troglodytes) are highly gregarious and form cooperative relationships with other males. In contrast, female social relationships vary within and between populations. Females in most East African populations, e.g., Gombe, Mahale, Kibale-Kanyawara, are less gregarious than males and spend most of their time alone or with only their dependent offspring. Researchers have attributed low female gregariousness to the high potential for feeding competition. I provide the first data on association patterns and agonistic interactions of female chimpanzees (Pan troglodytes schweinfurthii) from the unusually large Ngogo community, Kibale National Park, Uganda. Ngogo females were less gregarious than males, but spent a mean of 64% of their time in association with ≥1 other females and as much time in all-female parties as they did alone. Further, female dyads associated nonrandomly and they formed associative cliques. Association levels within cliques were similar to those among the relatively gregarious West African chimpanzee females at Taï (Pan troglodytes verus) and among bonobo (P. paniscus) females. Agonistic interfemale interactions were extremely rare, and monthly mean party size and the numbers of anestrous females per party do not correlate significantly with fruit availability. Thus, Ngogo females maintained relatively high levels of gregariousness, but avoided detrimental feeding competition by preferentially associating with a small subset of other community females.     

Allogrooming Behavior and Grooming Site Preferences in Captive Bonobos (Pan paniscus): Association with Female Dominance

I tested the utility of Seyfarth's (1977) model of rank-related attractiveness to explain the distribution of allogrooming behavior among captive bonobos (Pan paniscus). Adult female bonobos generally have high social status and may be dominant over males. As predicted by the model, I found that high-ranking adult females received most allogrooming within each of the four investigated groups. Among adult female-adult female dyads, however, allogrooming was not clearly associated with dominance rank. Contradictory to predictions of the model, the highest-ranking females were responsible for most displacements over allogrooming, and grooming competition is positively correlated with dominance rank. In the second part of this study, I investigated the social significance of allogrooming body site preferences. Bonobos direct significantly most allogrooming to the face of conspecifics, and high- and low-ranking individuals, as well as males and females, differ significantly in their preferences for certain allogrooming sites. Subordinates and males tended to avoid facial grooming and preferred the back and anogenital region, while high-ranking individuals and females directed most allogrooming to the face and head of grooming partners. Data from this study support the hypothesis that high-ranking females are the most attractive grooming partners within a female-centered bonobo society. Many other aspects of allogrooming behavior, however, are not consistent with the model of rank-related attractiveness.     

Association partners of young chimpanzees in the Mahale Mountains National Park,Tanzania

Association partners of young chimpanzees at the Mahale Mountains National Park were analyzed. Juvenile and adolescent chimpanzees associated frequently with their mothers, although mother-offspring association decreased as the offspring grew up. Males tended to leave their mothers and associate with adult males, while females remained frequently associating with their mothers in early adolescence. In late adolescence and young adulthood, males usually associated with adult males and cycling adult females. Females may transfer into neighboring unit-groups in this stage. Although an immigrant female tended to be alone when her estrous cycle stopped, she associated with many individuals, in particular with adult males, when she resumed cycling. Some orphans were observed to associate frequently with particular adults. The findings were discussed in relation to the unique characteristics of chimpanzee social system.     

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.     

Development of partner preferences in Japanese Macaques (Macaca fuscata): Effects of gender and kinship during the second year of life

Quantitative data are presented on the effects of subject sex, partner sex,and kinship on the social interactions of 18 juveniles of the Oregon troop of Japanese macaques (Macaca fuscata).Data on these subjects as infants were also used to detail maturational changes in partner sex preferences. Nine males and nine females, whose multiparous mothers represented a cross section of dominance ranks, were observed using a focal-animal technique. Juveniles of both sexes engaged in more proximity, contact, grooming, mounting, aggression, and social play with kin than with nonkin partners. They initiated less contact with females and more contact with males during their second year. They initiated more grooming and aggression during their second year than their first year, with females displaying a strong preference for grooming females and males specifically aggressing males more during the second year. Aggression was higher between same-sexed partners than between opposite-sexed partners. Males engaged in more social interactions with males during the second year than the first year of life. Males played more than females during both years. Males played more with males during the second year than the first year, and males played with males more than did females during the second year. We conclude that sex differences in behavioral frequencies become evident during the first year of life, and sex differences in partner preferences emerge during the second year of life.