New cases of inter-community infanticide by male chimpanzees at Ngogo,Kibale National Park,Uganda

Infanticide by males has been recorded in four chimpanzee populations, including that in Kibale National Park, Uganda. Some infanticidal attacks occur during inter-community aggression. The sexual selection hypothesis does not easily explain these attacks because they may not directly increase male mating opportunities. However, females in the attackers’ community may benefit by expanding their foraging ranges and thereby improving their reproductive success; thus infanticide may increase male reproductive success indirectly. We report two new cases of infanticide by male chimpanzees at Ngogo, Kibale National Park. Like two previous cases, these occurred during a boundary patrol and were almost certainly between-community infanticides. The patrolling males attacked despite the proximity of males from the victims’ presumed community. This probably explains why, unlike the earlier cases, they did not completely cannibalize their victims. Such attacks seem to be relatively common at Ngogo and infanticide may be an important source of infant mortality in neighboring communities. Our observations cannot resolve questions about the sexual selection hypothesis. However, they are consistent with the range expansion hypothesis: the infanticides occurred during a period of frequent encounters between communities associated with a mast fruiting event, and Ngogo community members greatly increased their use of areas near the attack site during another mast fruiting event one year later. Our observations contribute to growing evidence that lethal intergroup aggression is a common characteristic of wild chimpanzee populations.     

New Intercommunity Infanticides by the Chimpanzees of Ngogo,Kibale National Park,Uganda

Hostile intercommunity relations, including attacking and killing extra-community infants of both sexes have occurred at most wild chimpanzee sites. We describe three recent cases of intercommunity attacks on infants committed by members of the Ngogo chimpanzee community in Kibale National Park, Uganda. Two of the attacks resulted in confirmed infanticides while a third attack probably resulted in the infant's death. In common with previous accounts of chimpanzee infanticides, the attacks described here occurred during boundary patrols outside the Ngogo community's usual range, adult and adolescent males were the main participants, one infant was cannibalized after being killed, and the victims’ mothers did not accompany the attacking party back to the Ngogo range. However, the patrol parties during each infanticide were larger than before and included females from the Ngogo community. Our observations indirectly support both the range expansion and imbalance of power hypotheses, which address why and under which conditions chimpanzee intercommunity encounters lead to aggression. These cases of intercommunity infanticide add to the growing database of the phenomenon in wild chimpanzees.     

Demographic influences on the hunting behavior of chimpanzees

We investigated hunting in an unusually large community of wild chimpanzees at Ngogo in the Kibale National Park, Uganda. Aspects of predation were recorded with respect to the prey, the predators, and hunting episodes. During 23 months of observation, the Ngogo chimpanzees caught 128 prey items from four primate and three ungulate species. Chimpanzees preyed selectively on immature red colobus primarily during group hunts, with adult males making the majority of kills. Party size and composition were significant predictors of the probability that chimpanzees would hunt and of their success during attempts. Chimpanzees were more likely to hunt red colobus if party size and the number of male hunters were large; party size and the number of male hunters were also significantly larger in successful compared with unsuccessful hunts. The Ngogo chimpanzees did not appear to hunt cooperatively, but reciprocal meat-sharing typically took place after kills. Hunts occurred throughout the year, though there was some seasonality as displayed by periodic hunting binges. The extremely high success rate and large number of kills made per successful hunt are the two most striking aspects of predation by the Ngogo chimpanzees. We compare currently available observations of chimpanzee hunting behavior across study sites and conclude that the large size of the Ngogo community contributes to their extraordinary hunting success. Demographic differences between groups are likely to contribute to other patterns of interpopulation variation in chimpanzee predation. Am J Phys Anthropol 109:439–454, 1999. © 1999 Wiley-Liss, Inc.     

Grooming Between Male Chimpanzees at Ngogo,Kibale National Park. I. Partner Number and Diversity and Grooming Reciprocity

Allogrooming serves many social functions in primates. Grooming can help individuals to service social relationships generally, sometimes reciprocally, and may be particularly important in the development and maintenance of alliances. However, time constraints limit the number of partners with whom one individual can groom enough to maintain cooperative relationships. As a result, the size of its grooming network may reach an asymptote as the size of its group increases, and it may distribute its grooming less equally among potential partners. Chimpanzees live in multimale, fission-fusion communities; males are philopatric, and commonly associate and groom with each other. Males form within-community alliances that influence dominance rank and access to mates, and allies groom with each other regularly; males also cooperate in aggression between communities. The chimpanzee community at Ngogo, in Kibale National Park, Uganda, is unusually large and has more males than any other known community. Field data show that adult Ngogo males groomed far more with other adult males than with females or with adolescent males, in contrast to a previous report (Ghiglieri, 1984). Adolescent males groomed adults much more than the reverse; males groomed and were groomed by females about equally. Individual males groomed mostly with a small number of other males. On average, males at Ngogo had only slightly more male grooming partners overall and had the same number of important partners as those of males in a much smaller community in the Mahale National Park, Tanzania, and they distributed their grooming less equitably. These results fit those expected if limits on available grooming time cause males to have a loyalty problem as the number of potential grooming and alliance partners increases. Despite differences in the extent and equitability of their grooming networks, males at both Ngogo and Mahale showed reciprocity in grooming. Grooming reciprocity has been demonstrated for captive chimpanzee males, but the Ngogo findings are the first demonstrations of reciprocity in wild communities.     

Lethal intergroup aggression by chimpanzees in Kibale National Park, Uganda

Chimpanzees (Pan troglodytes) have hostile intergroup relations throughout most or all of their geographic range. Hostilities include aggressive encounters between members of neighboring communities during foraging and during patrols in which members of one community search for neighbors near territory boundaries. Attacks on neighbors involve coalitions of adult males, and are sometimes fatal. Targets include members of all age/sex classes, but the risk of lethal intergroup coalitionary aggression is highest for adult males and infants, and lowest for sexually swollen females. The best-supported adaptive explanation for such behavior is that fission-fusion sociality allows opportunities for low-cost attacks that, when successful, enhance the food supply for members of the attackers' community, improve survivorship, and increase female fertility. We add to the database on intergroup coalitionary aggression in chimpanzees by describing three fatal attacks on adult males, plus a fourth attack on an adult male and an attack on a juvenile that were almost certainly fatal. Observers saw four of these attacks and inferred the fifth from forensic and behavioral evidence. The attackers were males in two habituated, unprovisioned communities (Ngogo and Kanyawara) in Kibale National Park, Uganda. We also summarize data on other intercommunity attacks at Ngogo. Our observations are consistent with the "imbalance of power" hypothesis [Manson & Wrangham, Current Anthropology 32:369-390, 1991] and support the argument that lethal coalitionary intergroup aggression by male chimpanzees is part of an evolved behavioral strategy.     

Comparative Feeding Ecology of Two Communities of Chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) in Kibale National Park,Uganda

Several recent studies have documented considerable intraspecific and intrapopulation ecological variation in primates. However, we generally lack an understanding of how such variability may be linked to concomitant demographic variation among groups or populations of the same species, particularly in regard to large-bodied and wide-ranging species with high ecological flexibility, such as chimpanzees (Pan troglodytes). We compared the feeding ecology of chimpanzees inhabiting 2 sites in Kibale National Park, Uganda that differ 3-fold in chimpanzee density and support notably different plant communities. Chimpanzees at Ngogo, a site with the largest known chimpanzee community and unusually high chimpanzee density, spent a significantly lower percentage of time resting (and pregnant and lactating females spent more time feeding), incorporated higher percentages of ripe fruit in their diet, had lower dietary diversity values, and had shorter and less variable average patch residency times than did their counterparts at the nearby Kanyawara site, which supports a relatively low density of chimpanzees. In addition, feeding party size was significantly and positively related to feeding patch size at Ngogo, but not at Kanyawara. Together these findings aid in explaining the noted disparity in chimpanzee community size and density between Ngogo and Kanyawara by suggesting that the diet of Ngogo chimpanzees is of higher overall quality than that of Kanyawara chimpanzees. They also highlight the potentially profound influence of even small-scale habitat heterogeneity on the ecology of primates. Researchers must take such influences into account when attempting to draw conclusions about species- or population-level characteristics.     

Effects of male group size,parity, and cycle stage on female chimpanzee copulation rates at Ngogo,Kibale National Park,Uganda

Chimpanzees have complex and variable mating strategies, but most copulations occur when females with full sexual swellings are in parties with multiple males and mate with most or all of those males. Daily copulation rates for fully swollen females vary at different times of a female’s cycle, among females, and across communities and populations. Variation in female age, parity, and cycle stage underlie some of this variation, but possible demographic effects on copulation rates have not been systematically investigated. Demographic variation can affect many aspects of behavior and ecology, including the frequency and success of different mating tactics. Analysis of data from the unusually large chimpanzee community at Ngogo produces two results that are consistent with the hypothesis that demographic variation affects female copulation rates. Copulation rates were high compared with those reported from other research sites, where females had fewer potential mates available. Daily copulation rates of fully swollen females were also positively related to the number of males with whom they associated. Ngogo data also re-confirm results from other studies, of both wild and captive populations, showing that female copulation rates increase during periovulatory periods. This is consistent with the hypothesis that sexual swellings and extended receptivity and proceptivity help to protect females against infanticide by helping to ensure they mate with all potential sires. As at some other sites, parous females at Ngogo copulated at higher rates than nulliparous females. Possible effects of demography on sexual behavior should be considered in assessments of differences between chimpanzees and bonobos and of variation across chimpanzee populations.     

Grooming Between Male Chimpanzees at Ngogo,Kibale National Park. II. Influence of Male Rank and Possible Competition for Partners

Allogrooming contributes to the development and maintenance of social relationships, including those that involve alliances, in many primate species. Variation in relatedness, dominance rank, and other factors can produce variation in the value of others as grooming partners. Several models have been developed to account for variation in the distribution of grooming in relation to dominance ranks. These start from the premise that individuals are attracted to high-ranking partners, but time limits, direct competition, and prior grooming engagement between high-ranking individuals can constrain access to them. Sambrook et al. (1995) formalized some of these models and showed the importance of taking group size variation into account when assessing them. Chimpanzees form multimale communities in which males are the philopatric sex. Males commonly associate and groom with each other; they also form dominance hierarchies and form alliances that influence dominance ranks and mating success. Both male rank and the rank distance between partners are significantly correlated with the distribution of grooming between males in an extremely large chimpanzee community at Ngogo, Kibale National Park, Uganda, that has more males than any other known community. High-ranking males had more grooming partners than mid- or low-ranking males. Grooming predominantly went up the dominance hierarchy, but was also concentrated among males that were close in rank. Rank and rank distance apparently both affected grooming independently of reciprocity in grooming and independently of the frequency with which males associated in temporary parties. However, the data do not clearly indicate how constraints on access to partners might have operated. Published data from a smaller chimpanzee community at Mahale show no rank or rank distance effect on male grooming. These results and earlier, conflicting findings on the association between dominance rank and grooming in male chimpanzees indicate that variation in group size, i.e., the number of males per community, probably influences the strength of any such effects, as happens for grooming between females in several cercopithecine species. Data on coalitions at Ngogo support the argument that high-ranking males are valuable social partners, and similarity in strategies of alliance formation may influence the distribution of grooming.     

Demographic influences on the behavior of chimpanzees

Recent research has revealed substantial diversity in the behavior of wild chimpanzees. Understanding the sources of this variation has become a central focus of investigation. While genetic, ecological, and cultural factors are often invoked to explain behavioral variation in chimpanzees, the demographic context is sometimes overlooked as a contributing factor. Observations of chimpanzees at Ngogo, Kibale National Park, Uganda, reveal that the size and structure of the unit group or community can both facilitate and constrain the manifestation of behavior. With approximately 150 individuals, the Ngogo community is much larger than others that have been studied in the wild. We have taken advantage of the unusual demographic structure of this community to document new and intriguing patterns of chimpanzee behavior with respect to hunting, territoriality, and male social relationships. Chimpanzees at Ngogo hunt often and with a considerable degree of success. In addition, male chimpanzees there frequently patrol the boundary of their territory and engage in repeated bouts of lethal intergroup aggression. By forming two distinct subgroups, male chimpanzees at Ngogo also develop social bonds above the level of dyadic pairs. While the sheer number of chimpanzees contributes to differences in hunting, patrolling, mating, and subgrouping at Ngogo, the demographic situation may also constrain behavioral interactions. At Ngogo, male chimpanzees who are closely related genetically through the maternal line do not appear to affiliate or cooperate with each other. Demographic constraints may be responsible for this finding. In this paper, I use these examples to illustrate how the demographic context affects the possible range of behavioral options open to individuals and ultimately contributes to the explanation of behavioral diversity in chimpanzees.     

Suitable Habitats for Endangered Frugivorous Mammals: Small-Scale Comparison,Regeneration Forest and Chimpanzee Density in Kibale National Park,Uganda

Landscape patterns and chimpanzee (Pan troglodytes schweinfurthii) densities in Kibale National Park show important variation among communities that are geographically close to one another (from 1.5 to 5.1 chimpanzees/km²). Anthropogenic activities inside the park (past logging activities, current encroachment) and outside its limits (food and cash crops) may impact the amount and distribution of food resources for chimpanzees (frugivorous species) and their spatial distribution within the park. Spatial and temporal patterns of fruit availability were recorded over 18 months at Sebitoli (a site of intermediate chimpanzee density and higher anthropic pressure) with the aim of understanding the factors explaining chimpanzee density there, in comparison to results from two other sites, also in Kibale: Kanyawara (low chimpanzee density) and Ngogo (high density, and furthest from Sebitoli). Because of the post-logging regenerating status of the forest in Sebitoli and Kanyawara, smaller basal area (BA) of fruiting trees most widely consumed by the chimpanzees in Kanyawara and Sebitoli was expected compared to Ngogo (not logged commercially). Due to the distance between sites, spatial and temporal fruit abundance in Sebitoli was expected to be more similar to Kanyawara than to Ngogo. While species functional classes consumed by Sebitoli chimpanzees (foods eaten during periods of high or low fruit abundance) differ from the two other sites, Sebitoli is very similar to Kanyawara in terms of land-cover and consumed species. Among feeding trees, Ficus species are particularly important resources for chimpanzees at Sebitoli, where their basal area is higher than at Kanywara or Ngogo. Ficus species provided a relatively consistent supply of food for chimpanzees throughout the year, and we suggest that this could help to explain the unusually high density of chimpanzees in such a disturbed site.     

Intracommunity Coalitionary Killing of an Adult Male Chimpanzee at Ngogo,Kibale National Park,Uganda

Intercommunity coalitionary killing of adult and adolescent males has been documented in two chimpanzee communities in the wild, and it was strongly suspected in a third. It may increase survivorship for the attackers, their mates, and their offspring by reducing the combined strength of hostile neighbors and/or by increasing territory size and food availability, and it may help the attackers to attract mates. Lethal coalitionary attacks by males on other male members of their own communities would not provide these benefits and are not expected, given the importance of cooperation among male community members in contests for dominance rank and in both defense and offense against neighboring males. Nevertheless, intracommunity coalitionary killings associated with struggles for alpha rank occur in the wild and in captivity, and observers have seen serious gang attacks on maturing adolescent and young adult males at Mahale and Budongo: the victim in the Budongo case was killed (Fawcett and Muhumuza, 2000). I describe a lethal attack on a young adult male by a large coalition of males from his community at Ngogo, Kibale National Park, Uganda. The Ngogo community is the largest known for chimpanzees and has an unusually large number of males. The attack was not related to a struggle for alpha rank: the victim was low-ranking and the community had a well-established alpha at the time. However, the victim had risen substantially in the male hierarchy over the past few years and might have appeared threatening to many higher-ranking males. Simultaneously, he associated relatively little with most other adult males, had relatively few grooming partners and was not well integrated into the male grooming network, and had no influential allies. The combination of these social factors with the unusual demographic circumstances – which presumably meant that mating competition was relatively high and the cost of losing one male relatively low – might have triggered the attack.     

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.     

Diet of chimpanzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Park, Uganda, 1. Diet composition and diversity

Chimpanzees (Pan troglodytes) are ecologically flexible omnivores with broad diets comprising many plant and animal foods, although they mostly eat fruit (including figs). Like other ecologically flexible nonhuman primates (e.g., baboons, Papio spp.) with broad diets, their diets vary across habitats. Much data on diets come from short studies that may not capture the range of variation, however, and data are scant on variation within habitats and populations. We present data on diet composition and diversity for chimpanzees at Ngogo, in Kibale National Park, Uganda, collected over a 15-year period, with a focus on the plant components of the diet. We compare Ngogo data to those on chimpanzees at the nearby Kibale site of Kanyawara, on other chimpanzee populations, and on some other frugivorous-omnivorous primates. Results support the argument that chimpanzees are ripe fruit specialists: Ngogo chimpanzees ate a broad, mostly fruit-based diet, feeding time devoted to fruit varied positively with fruit availability, and diet diversity varied inversely with fruit availability. Comparison of Ngogo and Kanyawara shows much similarity, but also pronounced within-population dietary variation. Chimpanzees fed much more on leaves, and much less on pith and stems, at Ngogo. Figs accounted for somewhat less feeding time at Ngogo, but those of Ficus mucuso were quantitatively the most important food. This species is essentially absent at Kanayawara; its abundance and high productivity at Ngogo, along with much higher abundance of several other important food species, help explain why chimpanzee community size and population density are over three times higher at Ngogo. High inter-annual variation at Ngogo highlights the value of long-term data for documenting the extent of ecological variation among chimpanzee populations and understanding how such variation might affect population biology and social dynamics.     

General Gregariousness and Specific Social Preferences among Wild Chimpanzees

Wild chimpanzees form temporary parties that vary in size and composition. Previous studies have revealed considerable intraspecific variation in party compositions. We examined patterns of association among age, sex, and reproductive classes of chimpanzees at Ngogo in the Kibale National Park, Uganda. We employed a class-based association index and a randomization procedure to control for confounding factors and to test for differences between classes. Results indicate that males associated with other males significantly more than expected if all classes behaved equivalently, while females generally associated with individuals of the same sex less than expected. To interpret these patterns we used two additional indices that separate associations into two components: general gregariousness and preference for particular classes of associates. Males and estrous females were more gregarious than other classes, while anestrous females were less so. After controlling for general gregariousness, adult males as a class showed no specific preference for associating with each other. Anestrous females preferred each other as party members, and estrous females avoided each other. These results are consistent with previous findings that adult males are more gregarious than females. They diverge from the standard picture of chimpanzee society, however, by suggesting a mutual affinity among anestrous females, but not among adult males as a class.     

Phrase structure of wild chimpanzee pant hoots: Patterns of production and interpopulation variability

Recordings and behavioral observations of wild chimpanzees were made over a 2-year period in the Kibale National Park, Uganda (Kanyawara and Ngogo communities) to investigate patterns of acoustic variability in long-distance calls. The phrase structures of pant hoots, the species-typical loud call given predominantly by adult males, were analyzed. Analysis revealed that the build-up phrase was frequently absent from the pant hoots of Kibale chimpanzees. By contrast, analysis of published data on Gombe and Mahale chimpanzees (Tanzania) indicated that these animals typically included the build-up in their calls. These results were interpreted as evidence for phrase-level differences between populations in the acoustic morphology of this compound call. Data on age and sex differences in the context of production of pant hoots were analyzed, and their relevance to the possibility that aspects of pant hoot acoustic morphology are learned is discussed. Adult males initiated pant hoots more than subadult males, and adult males joined other pant hooting individuals with pant hoots more than subadult males did. It is suggested that younger males may pant hoot with specific adult males preferentially and that this may affect the development of their pant hoot acoustic morphologies. A peculiar pant hoot variant previously reported from Gombe, the whimper hoot, is described from Kibale. The production of this call by low-ranking individuals suggests that there are social constraints on pant hooting in the chimpanzee community. Ideas concerning the effect of social relationships on interpopulation variability in vocal signals are briefly discussed. © 1996 Wiley-Liss, Inc.     

Tool use in insect foraging by the chimpanzees of Ngogo, Kibale National Park, Uganda

Chimpanzee tool use for resource acquisition has been reported at numerous research sites. The chimpanzees of the Kibale Forest, western Uganda, have not previously been observed to use tools in foraging for insects. Here I report the first observation of tool use by the chimpanzees of the Ngogo community of Kibale National Park, Uganda, in insect foraging. Three adult females, one adolescent male, and one juvenile male were observed making and using tools to probe into a fallen dead tree to collect insect and wood pieces. I discuss the importance of this observation, and the behavioral similarities with chimpanzees from other sites.     

Energetic costs of territorial boundary patrols by wild chimpanzees

Chimpanzees are well known for their territorial behavior. Males who belong to the same community routinely patrol their territories, occasionally making deep incursions into those of their neighbors. Male chimpanzees may obtain several fitness benefits by participating in territorial boundary patrols, but patrolling is also likely to involve fitness costs. Patrollers risk injury or even death, and patrols may be energetically costly and may involve opportunity costs. Although territorial patrols have been reported at all long‐term chimpanzee study sites, quantitative data on their energetic costs have not previously been available. I evaluated the energy costs of patrolling for male chimpanzees at Ngogo, Kibale National Park, Uganda during 14 months of observation. In 29 patrols and matched control periods, I recorded the distances covered and time spent traveling and feeding by chimpanzees. I found that male chimpanzees covered longer distances, spent more time traveling, and spent less time feeding during patrols than during control periods. These results support the hypothesis that chimpanzees incur energetic costs while patrolling and suggest that ecological factors may constrain the ability of chimpanzees to patrol. Am. J. Primatol. 72:93–103, 2010. © 2009 Wiley‐Liss, Inc.     

Floristic heterogeneity at Ngogo,Kibale National Park,Uganda and possible implications for habitat use by chimpanzees (Pan troglodytes)

Tropical forest tree communities exhibit heterogeneity at multiple spatial scales, with important implications for animals relying on these resources. However, different organisms may perceive heterogeneity in the floristic community in very different ways. Here, we characterize the overall extent of heterogeneity in the floristic community at Ngogo, in Kibale National Park, Uganda. Also, using information from studies on the diet of chimpanzees (Pan troglodytes) inhabiting Ngogo, we quantify the extent to which the habitat is likely perceived as heterogeneous by this species. The forest as a whole is slightly more diverse than comparable sites in the Congo Basin, but this diversity is driven by a relatively high proportion of rare species represented by few individuals. From the perspective of chimpanzees, the forest is, unsurprisingly, even more heterogeneous. Species that provide fruit for chimpanzees during times of low overall fruit abundance and that display interindividual synchrony in fruiting were the most common chimpanzee resource in our sample, whereas species that provide fruit during times of low overall fruit abundance and that display asynchronous fruiting were the least common. We discuss the implications of the differences in density and distribution of various classes of resources for chimpanzee habitat use and foraging efficiency.     

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.     

Range use of the forest chimpanzees of Kibale: Implications for the understanding of chimpanzee social organization

We contrast the range use patterns of male and female chimpanzees (Pan troglodytes) based on repeated sightings over three years of 19 individuals in the Kanyawara community of Kibale Forest Reserve, Uganda. Depending on how home range size was calculated, male chimpanzees used an area that was 1.5 to 2 times greater than that of females. There was no difference between the sexes in whether their home ranges were used in a clumped or uniform fashion. However, males were more likely to be seen in boundary areas than females. These results are discussed in light of previously proposed models of chimpanzee social organization. It is concluded that the scenario in which females have smaller core areas within the defended home range of the males is most strongly supported by the range use patterns observed in Kibale chimpanzees. © 1993 Wiley-Liss, Inc.