Female Coalitions Against Male Aggression in Wild Chimpanzees of the Budongo Forest

In the wild, female chimpanzees (Pan troglodytes) are subject to male aggression that at times can be prolonged or particularly violent. There are no reports of cooperative retaliation to such aggression, a strategy observed in the congeneric Pan paniscus, from the wild despite >4 decades of detailed behavioral study across a number of populations and its occurrence among captive female chimpanzees. If the reports from captivity represent an inherent capacity, then the absence of similar reports from wild populations suggests that females may be able to form coalitions only under appropriate ecological and demographic conditions. During a study of male and female aggressive interactions among chimpanzees of the Budongo Forest, Uganda, wild adult female chimpanzees sometimes formed coalitions with one another to retaliate against male aggression. This may be possible because these females tend to be more gregarious than in other populations of East African chimpanzees, as other studies of the same population have suggested; the extent and variation of female chimpanzee social strategies may, therefore, need reconsideration. Further, my observations strengthen the argument that at least some of the differences between chimpanzees and bonobos may be more of degree than of kind.     

Probable Community Transfer of Parous Adult Female Chimpanzees in the Budongo Forest, Uganda

Female chimpanzees with dependent offspring generally avoid border areas of their community's home range because they risk aggression and infanticide from extracommunity males. Typically, only nulliparous females risk crossing the boundary areas to transfer between communities; while immigration of parous females occurs, it is extremely rare and dangerous for the females and their offspring. In the Budongo Forest, Uganda, where researchers have continuously studied the Sonso community since 1990, our field data provide strong indications that ≥5 adult females with offspring have immigrated into the community. If the interpretation is correct, then it has fundamental implications for our understanding of female chimpanzee social strategies and dispersal patterns. Further, the identification of such a large number of new individuals within a short time frame is remarkable and suggests a major event, such as the breakup of a neighboring community or major habitat disturbance. We explore the evidence that points to the events as immigrations and discuss the implications for understanding the chimpanzee social system.     

Decaying Raphia farinifera Palm Trees Provide a Source of Sodium for Wild Chimpanzees in the Budongo Forest,Uganda

For some years, chimpanzees have been observed eating the pith of decaying palm trees of Raphia farinifera in the Budongo Forest, Uganda. The reasons for doing this have until now been unknown. An analysis of the pith for mineral content showed high levels of sodium to be present in the samples. By contrast, lower levels were found in bark of other tree species, and also in leaf and fruit samples eaten by chimpanzees. The differences between the Raphia samples and the non-Raphia samples were highly significant (p<0.001). It is concluded that Raphia provides a rich and possibly essential source of sodium for the Budongo chimpanzees. Comparison of a chewed sample (wadge) of Raphia pith with a sample from the tree showed a clear reduction in sodium content in the chewed sample. Black and white colobus monkeys in Budongo Forest also feed on the pith of Raphia. At present, the survival of Raphia palms in Budongo Forest is threatened by the use of this tree by local tobacco farmers.     

The diet of chimpanzees in the Budongo Forest Reserve, Uganda

Chimpanzee diets are highly variable, but in all cases dominated by ripe fruit. Previous work has suggested that in the Budongo Forest Reserve, Uganda, ripe fruit remains available throughout the year, but dietary data for chimpanzees at this site remain sparse. This paper details the diet of a recently habituated community of Budongo chimpanzees living in the Sonso region. Chimpanzees of this community spent 64.5% of their feeding time eating fruit, and 19.7% eating arboreal leaves. Terrestrial herbaceous vegetation formed only a minor component of the diet. In these general dietary characteristics, Budongo chimpanzees show greater similarity to those of the Gombe National Park than they do to those in the geographically closer and floristically more similar Kibale Forest National Park. The diet was dominated by the fruit of four species and leaves of two species, although the composition of the diet varied from month to month, remaining diverse. Figs were consumed throughout much of the year, and in consequence, should perhaps be regarded as a staple, rather than fallback, food. A period of food scarcity was not apparent in this study and future comparative nutritional investigation will be required to determine whether these chimpanzees face a time of dietary hardship.     

Censusing chimpanzees in the Budongo Forest,Uganda

It is difficult to make valid comparisons of chimpanzee densities among sites because observers calculate them using different methods. We argue that nest count estimates of density are preferable to densities from home range estimates because of the problems of defining home range. There are many problems associated with nest count methods, some of which have not been addressed in previous studies. In 1992, we censused chimpanzees in the Budongo Forest using three methods;the standing crop nest count (SCNC), the marked nest count (MNC), and visual sightings of the animals (VS). Each method is based on standard line transect techniques. Of 96 nests monitored for decay rate,those constructed in the dry seasons decayed faster than those in the wet seasons. All- day follows of individual chimpanzees and observations of nesting chimpanzees at dusk showed that about 15.8% of night nests were reused,17.5% of the population did not build nests, and 18.8% of nests were first constructed as day nests. Given the variability in nest decay rates, we argue that MNC is a better method than SCNC because it avoids having to calculate decay rates.     

Social behavior of chimpanzees in the Budongo Forest,Uganda

The author studied the social organization and behavior of wild chimpanzees in the Budongo Forest, Uganda, between September, 1966, and March, 1967. A population of more than 50 chimpanzees became habituated to the author and was followed continuously, by means of individual identification, throughout the study period. The present article deals with the chimpanzee's social behavior, with special reference to social organization.The central organization which controls and maintains chimpanzee society cannot be rigid dominance rank order, since agonistic interactions among chimpanzees were not frequent. On the other hand, friendly or associative social behavior was frequently observed, some of which was unique in non-human primates. Greeting behavior, especially, and begging-and-food-sharing behavior show the existence of the chimpanzee's unique communicative system, which is backed by symbolic function. This kind of communicative system appears to have an important role in the maintenance of the unique social organization of chimpanzees, which has a loose regional integration, though the animals frequently gather into parties and later split up.Sexual activity was observed throughout the study period, but the peak was seen between December and February, during the dry season.The field work necessary for this study was carried out as a part of the Kyoto University Africa Primatological Expedition and was financed mainly by a Grant for Scientific Research from the Ministry of Education and partially by the Wenner-Gren Foundation for Anthropological Research and by the economic world of Japan.     

Social organization of chimpanzees in the Budongo Forest,Uganda

A population of wild chimpanzees in the Budongo Forest, Uganda, was studied between September, 1966, and March, 1967, by means of habituation and identification of each individual. This article deals with the grouping patterns, social units, and social organization of the chimpanzees in their natural habitat. Although parties of chimpanzees form groups of many patterns according to each particular situation and have no permanent membership, many chimpanzees live within a certain restricted area and gather to form parties consisting, in the main, of chimpanzees acquainted with each other. More than 50 individuals combined by social bonds in a loose regional population which is separated from other populations without recourse to geographical or physical barriers, though it maintains friendly contact with neighboring chimpanzee residents and with strangers. No particular individual group leader can be found, but any adult, especially a male, may act as a nucleus of each party.This study was carried out as a part of the Kyoto University Africa Primatological Expedition; the field work was financed mainly by a Grant for Scientific Research, Ministry of Education, and in part by the Wenner-Gren Foundation for Anthropological Research and by the economic world of Japan.     

Patterns of frugivory of the Budongo Forest chimpanzees, Uganda

Frugivory patterns of the chimpanzees in the Budongo Forest Reserve, Uganda were studied between June 2000 and August 2001. Chimpanzee feeding habitats, movement, group size and food eaten were assessed using focal and scan sampling. It was found that fruits were scarce during the dry season, when chimpanzees appeared and moved in large groups over long distances and raided farms at the forest edge. Chimpanzee movement out of the forest to forage was influenced by seasonal fluctuations in availability of preferred foods as some cultivated crops are perennial. Presence of chimpanzees in a specific feeding habitat was related to the availability of edible fruits both within and between months, suggesting that the presence of food may influence chimpanzee movement patterns. Therefore, a good understanding of patterns of frugivory is essential for making informed decisions about conservation of chimpanzees and other frugivores like birds and monkeys in Budongo as different forest habitats are under varying human pressure because of logging and other forms of utilization.     

Altruism in Forest Chimpanzees: The Case of Adoption

In recent years, extended altruism towards unrelated group members has been proposed to be a unique characteristic of human societies. Support for this proposal seemingly came from experimental studies on captive chimpanzees that showed that individuals were limited in the ways they shared or cooperated with others. This dichotomy between humans and chimpanzees was proposed to indicate an important difference between the two species, and one study concluded that “chimpanzees are indifferent to the welfare of unrelated group members”. In strong contrast with these captive studies, consistent observations of potentially altruistic behaviors in different populations of wild chimpanzees have been reported in such different domains as food sharing, regular use of coalitions, cooperative hunting and border patrolling. This begs the question of what socio-ecological factors favor the evolution of altruism. Here we report 18 cases of adoption, a highly costly behavior, of orphaned youngsters by group members in Taï forest chimpanzees. Half of the adoptions were done by males and remarkably only one of these proved to be the father. Such adoptions by adults can last for years and thus imply extensive care towards the orphans. These observations reveal that, under the appropriate socio-ecologic conditions, chimpanzees do care for the welfare of other unrelated group members and that altruism is more extensive in wild populations than was suggested by captive studies.     

The home range of the Sonso community of chimpanzees from the Budongo Forest, Uganda

Early studies of chimpanzees in the Budongo Forest, Uganda, reported only rough estimates for home range size. This study aimed to quantify home range size and habitat composition for chimpanzees of the Sonso community in this forest. Home range is the habitually used portion of the total area over which an animal travels, and for chimpanzees the home range is shared by the community. From data collected in 1994 and 1995, the home range for this community was estimated to cover an area of 6.78 km² using the minimum convex polygon analysis technique and 6.89 km² using fixed kernel analysis. Habitat was highly heterogeneous. The small home range and consequent high local population density (approximately 6.8 individuals km^?1) suggests that the Sonso region is highly productive in chimpanzee foods, possibly in contrast to other parts of the forest. This estimate of home range size is among the smallest reported for habituated chimpanzees, but comparisons with other sites are confounded by methodological and analytical differences. Future studies of chimpanzee ranging patterns should endeavour to use precise locations and the latest analytical techniques to ensure accurate estimation of range size and structure, and intersite comparability.     

Phenology of figs in Budongo Forest Uganda and its importance for the chimpanzee diet

This paper reports on the phenological patterns of figs in Budongo Forest, Uganda, and how it relates to chimpanzee food availability in different seasons. In addition, we analysed the dung of chimpanzees to understand the composition of fruits in their diet. The aim of our study was to assess Ficus phenology and how it affects chimpanzee diet. Fifteen species of figs were monitored for fruit (syconium) and leaf phenology between June 2000 and 2001. Ficus fruit production varied significantly between and within species, and also with tree trunk and crown diameters. Fig fruit production was asynchronous and individual fig trees produced crops from one to five times in a year. In addition to fruits, chimpanzees fed on young leaves of some Ficus species. Shedding of old Ficus leaves coincided with the dry season, followed by appearance of young leaves. The dry season in Budongo is a period of general fruit scarcity. The combination of fig fruits and young leaves make up the most important food in the diet of chimpanzees. From the chimpanzee dung, more than 78% of seeds comprised fig ‘seeds’ (nutlets) and the rest of the diaspores were from other tree species. Our findings suggest that chimpanzees disperse large number of diaspores in their dung, thereby serving as important agents of natural forest regeneration.