Social behavior and relationship among wild chimpanzees of the Mahali mountains

Patterns of social behavior of the chimpanzees of the Mahali Mts. are listed briefly in this paper, where behavior absent in the other subhuman primates is especially focussed upon. Some are more analogous to those of mankind than to those of the other subhuman primates. This characteristic social behavior is more or less related to the peculiar social organization of wild chimpanzees. Differences of social behavior between two local populations (i.e., of the Mahali Mts. and the Gombe Stream Reserve) have not been recognized; namely, dialect differentiation does not seem to exist.     

Drinking tools of wild chimpanzees at Bossou

Use of drinking tools by wild chimpanzees (Pan troglodytes) and the context in which the tools were used were studied at Bossou, Republic of Guinea, West Africa. During the middle to late dry season and early wet season liquids are available occasionally in the holes of trees. Chimpanzees drank water or sap using a leaf (or fiber) as a sponge or spoon. When the chimpanzees were on the ground, they tended to use one of a few kinds of soft, hairless leaves, if they were available nearby. Females, particularly juveniles and adolescents, were thought to be the main users of the drinking tool. In a few episodes, a tool set was used to procure liquid. Once a chimpanzee used a stick to push a leaf sponge into a water hole and to pull it out from the hole. In addition, three chimpanzees used a pestle to squeeze sap from an oil-palm tree before using a fiber sponge. © 1995 Wiley-Liss, Inc.     

Opportunistic and restrictive matings among wild chimpanzees in the Mahale Mountains,Tanzania

The mating patterns of free-ranging chimpanzees (Pan troglodytes) of the Mahale Mountains, Tanzania, were studied. Opportunistic mating (non-competitive and temporary mating) was frequently observed in a large-sized unit-group, among young, low-ranking males, and among young, newcomer, non-ovulating females. Restrictive mating (a continuous sexual relationship between a particular pair which includes possessiveness and consortship) was frequently observed in a small-sized unit-group, among middle- and old-aged, high-ranking males, and among old, resident, ovulating females. Relations between those characteristics, such as group size and composition, ages of the individuals of both sexes, female estrous stages, and life history, and the 2 mating patterns are discussed.     

Reconciliation and post-conflict third-party affiliation among wild chimpanzees in the Mahale Mountains,Tanzania

This study investigated post-conflict (PC) behavior among wild chimpanzees (Pan troglodytes) of the M-group in the Mahale Mountains, Tanzania, and examined what types of behavior characterize the PC situation in this group, and the factors that influence the occurrence of PC affiliation between opponents soon after the end of an aggressive conflict (i.e., reconciliation). We found that the opponents affiliated selectively soon after the end of aggression, suggesting that reconciliation occurred in this group. The mean individual corrected conciliatory tendency (CCT) (Veenema et al. 1994 in Behav Proc 31:29–38) was 14.4%, which is similar to or lower than frequencies observed in studies of captive and wild chimpanzees. The valuable relationship hypothesis predicts that the CCT is higher among individuals who share valuable relationships (e.g., males or affiliative dyads) than among individuals who do not (e.g., females or less-associative dyads). However, the analysis based on data for aggression between unrelated individuals (including one incident between an adult and non-adult) and aggression between unrelated adults, did not uncover this difference. Affiliation by a previously uninvolved individual with the victim (consolation) and with the aggressor (appeasement) occurred more frequently following aggression than in the control condition. The results are compared with previous studies of captive and wild chimpanzees.     

Mortality rates among wild chimpanzees

In order to compare evolved human and chimpanzees' life histories we present a synthetic life table for free-living chimpanzees, derived from data collected in five study populations (Gombe, Ta?, Kibale, Mahale, Bossou). The combined data from all populations represent 3711 chimpanzee years at risk and 278 deaths. Males show higher mortality than females and data suggest some inter-site variation in mortality. Despite this variation, however, wild chimpanzees generally have a life expectancy at birth of less than 15 years and mean adult lifespan (after sexual maturity) is only about 15 years. This is considerably lower survival than that reported for chimpanzees in zoos or captive breeding colonies, or that measured among modern human hunter-gatherers. The low mortality rate of human foragers relative to chimpanzees in the early adult years may partially explain why humans have evolved to senesce later than chimpanzees, and have a longer juvenile period.     

Tool use by wild chimpanzees in feeding upon driver ants

The use of stick tools by wild chimpanzees (Pan troglodytes) to feed upon driver ants (Dorylus (Anomma) nigricans) is described. Observations of this ant dipping were made over five years in the Gombe National Park in western Tanzania. Chimpanzees find the nomadic ant colonies visually, often re-visiting the subterranean nest sites until the ants move on. The nest is opened manually and tools are made from green woody vegetation. The ant dipping sequence is intricate and efficient. The chimpanzee predator uses several positioning strategies to minimise the ants' massed defensive tactics. Average intakes of 17–20 g/feeding session are estimated. It is concluded that ants may constitute a significant chimpanzee dietary component and that the dipping tools and techniques are relatively stereotyped in form.     

Predatory behavior among wild chimpanzees of the mahale mountains

Eighteen predatory and two cannibalistic episodes have been observed among wild chimpanzees of the Mahale Mountains. The prey consists mainly of juveniles of medium-sized mammals, six species of which were recorded as prey fauna for the first time in this study. Predatory behavior of Mahale chimpanzees seems more opportunistic and primitive than has ever been observed in the Gombe National Park in terms of capture technique, size of prey, co-operation in hunting, sexual difference in capture frequency, degree of consumption and extent of meat sharing. Cannibalism is an inter-unit-group phenomenon and may be an extension of infanticide, which may function in changing “mother” into “female.” This study was financed mainly by the Overseas Scientific Research Fund of the Ministry of Education, Japan and partly by the Japan International Co-operation Agency and the Ministry of Natural Resources and Tourism, Tanzania.     

Dynamics of the temporal structures of playing clusters and cliques among wild chimpanzees in Mahale Mountains National Park

The overall structure and temporally changing configuration of members of social play among the wild chimpanzees (Pan troglodytes schweinfurthii) of Mahale Mountains National Park, Tanzania, were described on both the microscopic ‘clique’ levels, conceptualized as directly connected configurations through social play behavior, and macroscopic ‘cluster’ levels, conceptualized as indirectly connected gatherings of members of adjacent multiple cliques at the same time and space. Most playing clusters started as dyads. Although the cumulative number of participants increased, each clique size remained at 2 for most of the observational units. Dyadic cliques were more stable and lasted longer than larger cliques. Of all clusters’ playing fields, 64.7 % had maximum diameters of 3 m. In summary, chimpanzees played stably in dyadic pairs in most of the time. As time passed, other chimpanzees often joined in the playing groups to form large polyadic clusters. Even when all chimpanzees in a cluster played socially at the same time, they normally did so in separate dyadic pairs, forming multiple dyadic cliques simultaneously in a small space. These social play dynamics may be explained assuming a hypothesis based on a balance model among socially playing chimpanzees, as the balanced cliques are limited only to those in which all the existing pairs form the mutual dyads, and they tend to avoid unbalanced and maintain balanced relationships during social play. As a result, larger cliques were difficult to maintain for long periods and tended to transition into dyadic mutual cliques. Thus, Heider’s balance theory can be one of the possible theories to explain not only human social phenomena, but also the proximate mechanism of the structure and the temporal change of social play among wild chimpanzees. Although both mutual and transitive relationships are known to be balanced in various human networks, only mutual relationships among socially playing chimpanzees were balanced.     

Aggression toward large carnivores by wild chimpanzees of Mahale Mountains National Park, Tanzania

In the Mahale Mountains National Park of Tanzania, a group of about 33 chimpanzees were observed to surround a leopard den containing a mother and at least one cub and to drag out and kill the cub. This is the first report of chimpanzees or any other primate species killing their potential predator's offspring. The incident suggests that chimpanzees, without any weapons, can manage to defend themselves against a carnivore of at least up to leopard size, and implicates how the early hominids may have reacted against their potential predators.     

One observed case of temporary adoption of an infant by unrelated nulliparous females among wild chimpanzees in the Mahale Mountains,Tanzania

A 2.4-year-old male chimpanzee in the Mahale Mountains, Tanzania became separated from his mother probably because of contagion of the flu-like sickness. He was adopted alternately by two unrelated immigrant females for at least six days before he was reunited with his mother. This episode is described and analyzed in relation to the babysitter relationships among female chimpanzees. The adaptive significance of the babysitting behavior of nulliparous females is discussed.     

Tool use and tool making in wild chimpanzees

Reported incidences of tool use and tool making for three wild chimpanzee populations increase from Mahale (12 and 3 types of use and making, respectively), Gombe (16 and 3) to Ta? (19 and 6). Sticks are commonly used and prepared at all three sites. However, Ta? chimpanzees seem to perform more modifications on the material before using it. They are also the only chimpanzees seen to pound objects with tools and to combine two different tool uses to get access to one food item. Tool making is the rule for abundant material (grass, twigs), but appears to be rarer for scarce, hard material (clubs, stones). Factors involved in the acquisition and the benefit of tool use are discussed along with factors affecting the frequency and complexity of tool making in chimpanzees.     

Hand preference and tool use in wild chimpanzees

The hand preference of chimpanzees in their natural habitat was studied at Bossou, Republic of Guinea, West Africa. The quantitative difference in left/right hand use was small in food picking and carrying. In contrast, the chimpanzees employed either the right or left hand in nutcracking behavior using a pair of stones. All adults and many adolescents and juveniles utilized one hand exclusively for holding a hammer stone. Left hand preference was more prevalent among adults. However, when adolescents and juveniles were included, there was no significant bias in the ratio of left/right handers. Nut-cracking behavior requires long-term learning of the fine manipulation of stones and nuts by both hands. Each hand has a separate role, and the hands work together in nut cracking. The differential and complementary use of both hands may be a prime factor promoting exclusive hand preference in chimpanzees comparable to that of humans.     

Deactivation of snares by wild chimpanzees

Snare injuries to chimpanzees (Pan troglodytes) have been reported at many study sites across Africa, and in some cases cause the death of the ensnared animal. However, very few snare injuries have been reported concerning the chimpanzees of Bossou, Guinea. The rarity of snare injuries in this study group warrants further consideration, given the exceptionally close proximity of the Bossou chimpanzees to human settlements and the widespread practice of snare hunting in the area. Herein we report a total of six observations of chimpanzees attempting to break and deactivate snares, successfully doing so on two of these occasions. We observed the behavior in 5 males, ranging in age from juveniles to adults. We argue that such active responses to snares must be contributing to the rarity of injuries in this group. Based on our observations, we suggest that the behavior has transmitted down the group. Our research team at Bossou continues to remove snares from the forest, but the threat of ensnarement still remains. We discuss potential ways to achieve a good balance between human subsistence activities and the conservation of chimpanzees at Bossou, which will increasingly be an area of great concern in the future.     

Tool use by wild chimpanzees (Pan troglodytes) to obtain termites (Macrotermes herus) in the Mahale Mountains,Tanzania

Wild chimpanzees (Pan troglodytes) used probes made of vegetation to fish for termites (Macrotermes herus) in the Mahale Mountains of western Tanzania. Data on both the artefacts and behavior associated with their use were recorded over a 5-month period. The chimpanzees of one unit group, B Group, were seen four times to use the tools to extract the insects from their mounds. A sample of 290 tools was collected and analyzed in terms of age, seasonality, length, width, class of raw materials, species, methods of making, extent of use, and damage incurred through use. There were differences across groups in tool use in feeding on termites; some could be related to biotic factors, but others appeared to be cultural differences.     

New data on the use of stone tools by chimpanzees in Guinea and Liberia

Two types of use of “hammers” for cracking nuts by wild-living chimpanzees have been distinguished: (1) Relatively small stones are used by the chimpanzee community at Bossou in Guinea to crack the nuts of oil palms growing on abandoned farmland, while no nuts of wild tree species are cracked. (2) Larger hammer stones (and, at some sites, wooden clubs) are used in a more sophisticated manner to crack the nuts of wild trees, but not of oil palms, in an area ranging from south-east Sierra Leone through Liberia to the south-west of the Ivory Coast. The first author (1986) has proposed that Type I has been copied by the chimpanzees, under pressure of food shortage, from the local human population. New data now indicate that, at Bossou, while habitat deterioration has continued, the number of hammer and anvil stones per utilized oil palm tree has approximately tripled in the last six years. The quantity of food obtained from oil palm nut kernels, however, amounts to only a few percent of the total diet. For the rest these apes depend to a large extent on many other agricultural products cultivated at Bossou which they are allowed freely to consume, including even cassava (manioc) roots and sweet potatoes dug by them from the ground. Some factors determining the chosen size of hammers were analyzed. Two abnormal hammers were found whose wear suggested a tentative, human-like manner of use. No evidence has been found to indicate the use of stone tools by chimpanzees in the adjoining chimpanzee-inhabited areas around the range of the Bossou community. Type II stone tool use was found, however, in a primary forest on a mountain≈13 km west of Bossou. This is especially intriguing because the site is separated by a wide belt of drier rain forest from the belt of very humid rain forest in the south where all the other known Type II sites are located. More research on the geographical distribution of the use of stone tools by chimpanzees and on the underlying ecological factors is recommended.     

Lions eat chimpanzees: The first evidence of predation by lions on wild chimpanzees

No report has confirmed predation on wild chimpanzees by lions. Recently, at least two lions visited the Mahale Mountains National Park, Tanzania. Chimpanzee remains, including hairs, bones, and a tooth, were found in four samples of lion feces between June and September 1989. The discovery of each fecal sample, widely separated in time, combined with analyses of daily attendance records and fecal contents, suggests that at least four chimpanzees, including two adult females, an old adolescent male, and an early adolescent male, were eaten. Observed responses of chimpanzees to lions included alarm calls, whimpers, climbing into trees, and silence. Predation by lions has been neglected as a factor shaping the social structure of chimpanzees. Nevertheless, the large party size that has often been observed in savanna-living chimpanzees, which are sympatric with lions, might be an anti-predator response of this species. © 1993 Wiley-Liss, Inc.     

A case report of a novel type of stick use by wild chimpanzees

We have found evidence that wild chimpanzees used stout sticks to dig into one end of a decayed fallen trunk from the side and a long stick with a frayed end to dig into or brush its stump, in the Moukalaba Reserve, Gabon. This type of stick use by wild chimpanzees has not been recorded in any habitat. This finding should contribute to future studies and discussions of variations in tool use and cultural processes among wild chimpanzees. Electronic Publication     

One observed case of cannibalism among wild chimpanzees of the Mahale mountains

This paper presents a detailed report on the behavior of cannibalism observed in a group of wild chimpanzees of Western Tanzania for two days of January 13 and 14, 1977. This case differs from the other ones observed so far in that the adult males killed and ate an infant of their own group. Much interesting behavior was observed in the mother whose infant was eaten and the high-ranking males who killed and ate it.