New observations on the tool-behavior of chimpanzees from Mt. Assirik (Senegal,West Africa)

The tool-behavior of chimpanzees were studied at the Mt. Assirik, in the Parc National du Niokolo-Koba, Senegal from December 1982 to June 1983 and from February 1985 to January 1986. Five leaf-stalks to obtain termites ofMacrotermes subhyalinus species and four sticks to obtain honey (this is the first report on the use of tools to obtain honey by wild chimpanzees in far western Africa) were found; we also report the first evidence of tools probably used as hammers to break open hard-shelled fruits ofAdansonia digitata. We conclude that Mt. Assirik chimpanzees provide evidence of a certain cultural behavior related to the use of stones.     

Interspecific interactions between wild pygmy chimpanzees (Pan paniscus) and red colobus (Colobus badius)

Interspecific interactions accompanied by physical contacts between wild pygmy chimpanzees (Pan paniscus) and red colobus (Colobus badius) were observed on three occasions at Wamba, Republic of Zaire. In all cases, the red colobus initiated the interactions by approaching the pygmy chimpanzees. Most of the pygmy chimpanzees, which were within 5 m of the red colobus, were juveniles or infants but the adult male pygmy chimpanzees never showed any interest in the red colobus. The red colobus groomed the chimpanzees in two cases, but the latter never groomed the former. No true aggressive interactions were observed between the two species. The lack of any evidence of hunting of red colobus through longitudinal studies of the pygmy chimpanzees of Wamba, together with the present observations, suggests that red colobus are probably not targets of hunting by the pygmy chimpanzees.     

Do bonobos copulate more frequently and promiscuously than chimpanzees?

The copulatory activities of bonobos (Pan paniscus) of Wamba, Zaire, were compared with those of chimpanzees (P. troglodytes schweinfurthii) of Mahale, Tanzania. The copulation rates of adult male bonobos were equal to or lower than those of adult male chimpanzees. The copulation rates of adult female bonobos were approximately equal to those of adult female chimpanzees who were in maximal genital swelling, but it should be much higher than those of the adult female chimpanzees throughout the birth interval. The copulation rates of adolescent male bonobos were lower than those of adolescent male chimpanzees, whereas the copulation rates of adolescent female bonobos were much higher than those of adolescent female chimpanzees. It was suggested that the bonobos of Wamba did not copulate more promiscuously than did the chimpanzees of Mahale. The female bonobos may show “receptivity”, whereas female chimpanzees may show rather “proceptivity”.     

Behavioral Ecology of Sympatric Chimpanzees and Gorillas in Bwindi Impenetrable National Park,Uganda: Diet

Via a field study of chimpanzees (Pan troglodytes schweinfurthii) and gorillas (Gorilla gorilla beringei) in Bwindi Impenetrable National Park, Uganda, we found that their diets are seasonally similar, but diverge during lean seasons. Bwindi chimpanzees fed heavily on fruits of Ficus sp., which were largely ignored by the gorillas. Bwindi gorilla diet was overall more folivorous than chimpanzee diet, but was markedly more frugivorous than that of gorillas in the nearby Virunga Volcanoes. During 4 mo of the year Bwindi gorilla diet included more food species than that of the chimpanzees. Three factors in particular—seasonal consumption of fibrous foods by gorillas, interspecific differences in preferred fruit species, and meat consumption by chimpanzees—contributed to dietary divergence between the two species. When feeding on fruits, gorillas ate Myrianthus holstii more frequently than chimpanzees did, while chimpanzees included more figs in their annual diet. Chimpanzee diet included meat of duikers and monkeys; gorilla frequently consumed decaying wood.     

Comparison of behavioral sequence of copulation between chimpanzees and bonobos

We compared sex differences in behaviors leading to copulation of chimpanzees (Pan troglodytes) in the Kalinzu Forest, Uganda with those of bonobos (Pan paniscus) at Wamba, D.R. Congo, using the same definition. Female chimpanzees were more likely to initiate copulation than female bonobos. While most of copulations (96%) were initiated by males in bonobos, among chimpanzees only 63% of copulations were initiated by males. Female bonobos initiated an interaction leading to copulation when males approached them within a short distance. On the other hand, both male and female chimpanzees initiated behavior at a longer distance. Higher proceptivity and a higher copulation rate during the maximal swelling period of female chimpanzees might suggest that they gain greater benefits from a high frequency of copulations than do female bonobos.     

Group Size and Composition of Colobus guereza in Kyambura Gorge, Southwest Uganda, in Relation to Chimpanzee Activity

We documented and assessed the influence of chimpanzee activity on group size and composition of Colobus guereza in Kyambura Gorge, southwest Uganda, from July to September, 1994 and in February and March 1996. The population density of colobus is very high: 347 individuals per km ². Density differed outside activity centers of chimpanzees (525 individuals per km ² ) and within the centers (186 per km ² ). We identified a total of 24 colobus groups, ranging between 3 and 13 individuals. Of the 24 groups, 22 were one-male groups. Groups were smaller and the percentage of subadults and juveniles was lower in groups within chimpanzee activity centers. Estimates of home ranges are between 1.7 and 6.2 ha, but neighboring home ranges overlapped 80%. Only a territory of approximately 0.5 ha was defended by the alpha male.     

Behavioral diversity in populations of free-ranging chimpanzees in Africa: is it culture?

Behavioral differences exist among populations of wild chimpanzees (Pan troglodytes, Hominoidea) across Africa. Deciding if these are cross-cultural differences demands careful interpretation and rigorous scrutiny. Even standard patterns like social grooming may show variation in detail between neighboring populations. Some patterns of tool use, such as using hammers to crack nuts, also vary across far western Africa. At least 37 populations of free-ranging chimpanzees show tool use, and 14 show at least one habitual pattern of tool use. Regional differences exist among the three subspecies or geographical races of chimpanzees. Convincing empirical demonstration of social customs and traditions in non-human species is problematical, and even something so simple in principle as innovation is hard to show in practice. However, culture need not depend on imitation, teaching, or language, either in humans or in other species.     

Investigation of the RH locus in gorillas and chimpanzees

The human Rh blood-group system is encoded by two homologous genes,RhD andRhCE. TheRH genes in gorillas and chimpanzees were investigated to delineate the phylogeny of the humanRH genes. Southern blot analysis with an exon 7-specific probe suggested that gorillas have more than twoRH genes, as has recently been reported for chimpanzees. Exon 7 was well conserved between humans, gorillas, and chimpanzees, although the exon 7 nucleotide sequences from gorillas were more similar to the humanD gene, whereas the nucleotide sequences of this exon in chimpanzees were more similar to the humanCE gene. The intron between exon 4 and exon 5 is polymorphic and can be used to distinguish the humanD gene from theCE gene. Nucleotide sequencing revealed that the basis for the intron polymorphism is anAlu element inCE which is not present in theD gene. Examination of gorilla and chimpanzee genomic DNA for this intron polymorphism demonstrated that theD intron was present in all the chimpanzees and in all but one gorilla. TheCE intron was found in three of six gorillas, but in none of the seven chimpanzees. Sequence data suggested that theAlu element might have previously been present in the chimpanzeeRH genes but was eliminated by excision or recombination. Conservation of theRhD gene was also apparent from the complete identity between the 3′-noncoding region of the human D cDNA and a gorilla genomic clone, including anAlu element which is present in both species. The data suggest that at least twoRH genes were present in a common ancestor of humans, chimpanzees, and gorillas, and that additionalRH gene duplication has taken place in gorillas and chimpanzees. TheRhCE gene appears to have diverged more thanRhD among primates. In addition, theRhD gene deletion associated with the Rh-negative phenotype in humans seems to have occurred after speciation. Correspondence to: C.M. Westhoff     

Predation on mammals by chimpanzees in the montane forest of Kahuzi, Zaire

The rate of predation on mammals by chimpanzees was determined from carcasses and from fecal specimens found on fresh trails during a 16-month period in the montane forest of Kahuzi-Biega National Park, Zaire. A unit-group of semi-habituated chimpanzees, composed of 22 – 23 individuals including 8 adult or adolescent males, appeared to kill about 18 – 30 mammalian prey (16 – 28Cercopithecus monkeys) per year, if the multiple kills by chimpanzees were not considered. A juvenile l'Hoest's monkey was recorded for the first time as the prey of chimpanzees in this study. Predation occurred in the late dry and the early rainy seasons, when the diversity of ripe fruits was the highest during the year. The Kahuzi chimpanzees tended to kill mammals less frequently but to killCercopithecus monkeys more frequently than chimpanzees in other habitats. The absence of red colobus monkeys, which are the most frequent prey in Gombe, Mahale, and Tai, might be responsible for the low predation rate. However, the estimated rate of predation onCercopithecus monkeys is the highest record among various chimpanzee habitats. At least 11 – 18% of theCercopithecus population seemed to be lost annually as a result of being killed by chimpanzees. Chimpanzees may be the most important predators on these monkeys in the absence of leopards at Kahuzi. The examination of fecal samples and carcasses suggested that adult (probably male) or adolescent chimpanzees tended to eat juvenile or subadult monkeys most frequently, as is also seen for chimpanzees in Gombe, Mahale, and Tai.     

A note on the chimpanzee ecology of the Ugalla Area,Tanzania

Results of a nine-day survey of the chimpanzee habitat in Ugalla area of western Tanzania in 1975 is reported. The most dominant vegetation of the area wasJulbernardia globiflora woodland. Evidence of chimpanzees, both indirect (such as beds, faeces, and food remnants) and direct (calls and actual observation), suggests that chimpanzees are sparsely distributed throughout the area, but that they disproportionately useBrachystegia bussei woodland along escarpments andCynometra-Albizzia riverrine forests. The hilly country of Sisegwa/Mnyangwa was the only area where chimpanzees appeared to be comparatively abundant. Mammal hair found in faeces indicates mammal-predation by the chimpanzees in this area. Hard-shelled fruits ofStrychnos were apparently bitten open without the use of extra-somatic objects. The food repertoire included fruits of the generaAzanza, Canthium, Cordia, andTamarindus.     

Natural history of <Emphasis Type="Italic">Camponotus</Emphasis> ant-fishing by the M group chimpanzees at the Mahale Mountains National Park,Tanzania

The aim of this study was to provide basic data on ant-fishing behavior among the M group chimpanzees at the Mahale Mountains National Park, Tanzania. Ant-fishing is a type of tool-using behavior that has been exhibited by Mahale chimpanzees when feeding upon arboreal carpenter ants (Camponotus spp.) since the 1970s, and is now regarded as a candidate of wild chimpanzee culture. Herein, I describe in detail the features of ant-fishing shown by the Mahale M group chimpanzees: (1) 2 species of Camponotus ants (Camponotus sp. (chrysurus-complex) [C. sp.1] and C. brutus) were identified as the target species of ant-fishing, and C. sp.1 was selected intensively as the main target; (2) 24 species (92 individuals) of trees were identified as ant-fishing sites–these were widely distributed throughout the western/lowland region of the M group’s home range, and the top 5 species were used more frequently; (3) the efficiency of ant-fishing was influenced not only by the site choice or the skillfulness of the chimpanzees, but inevitably by the condition of the ants; (4) the estimated nutritional intake from ant-fishing was apparently negligible; (5) most of the M group members (50/60 individuals) older than 3 years of age successfully used tools to fish for ants; and (6) female chimpanzees engaged in ant-fishing more frequently and for longer periods than males did. Further, I compared the features of ant-fishing exhibited by the Mahale M group chimpanzees with those exhibited by the former K group at Mahale and by other populations of wild chimpanzees.     

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.     

Comparing infant and juvenile behavior in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes): a preliminary study

The dichotomy between the two Pan species, the bonobo (Pan paniscus) and chimpanzee (Pan troglodytes) has been strongly emphasized until very recently. Given that most studies were primarily based on adult individuals, we shifted the “continuity versus discontinuity” discussion to the infant and juvenile stage. Our aim was to test quantitatively, some conflicting statements made in literature considering species differences between immature bonobos and chimpanzees. On one hand it is suggested that infant bonobos show retardation in motor and social development when compared with chimpanzees. Additionally it is expected that the weaning process is more traumatic to chimpanzee than bonobo infants. But on the other hand the development of behaviors is expected to be very similar in both species. We observed eight mother–infant pairs of each species in several European zoos. Our preliminary research partially confirms that immature chimpanzees seem spatially more independent, spending more time at a larger distance from their mother than immature bonobos. However, the other data do not seem to support the hypothesis that bonobo infants show retardation of motor or social development. The development of solitary play, environmental exploration, social play, non-copulatory mounts and aggressive interactions do not differ between the species. Bonobo infants in general even groom other group members more than chimpanzee infants. We also found that older bonobo infants have more nipple contact than same aged chimpanzees and that the weaning process seems to end later for bonobos than for immature chimpanzee. Additionally, although immature bonobos show in general more signs of distress, our data suggest that the weaning period itself is more traumatic for chimpanzees.     

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.     

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.     

A note on the ecology of the red-colobus monkeys (Colobus badius tephrosceles) living in the Mahali Mountains

Red-colobus monkeys of the Mahali Mountains utilize many vegetation types; although their main habitat is composed of forest vegetation (gallery forest and montane forest), they sometimes penetrate into even theMiombo woodland.Red-colobus monkeys mainly live on leaves of trees, but sometimes eat flowers, fruits, and barks. Most of the plant leaves which the chimpanzees eat have proved to be delicacies to colobus monkeys also. Colobus monkeys live in compact bisexual groups, ranging from 30 to 50 in size. No solitary male has so far been observed.A colobus group living in the gallery forest moved 100 to 200 meters daily within its welldocumented home range. The home ranges of groups do not seem to overlap each other extensively.Their natural enemies are leopards, birds of prey and probably chimpanzees. The Tongwe inhabitants avail themselves of the colobus coat as part of their native bellows (mufuba) and some pagan inhabitants kill them for their meat.The research was financed by the Scientific Research Fund of the Ministry of Education and in part by the Wenner-Gren Foundation. This report is a part of the results of the Kyoto University Africa Primatological Expedition.     

Meat eating and ant dipping by wild chimpanzees in Sierra Leone

This report presents the first records of meat-eating and ant dipping by wild chimpanzees,Pan troglodytes, from Sierra Leone. The study was conducted in the proposed Outamba-Kilimi National Park, Northern Sierra Leone. Measurements of tools used to dip for driver ants, are compared with those from four other study sites in Africa. The results reveal some fundamentally common characteristics. From both faecal analysis and direct observation, evidence was found that the chimpanzees eat meat. These recordings indicate a varied choice of prey and add new species to those preyed upon by wild chimpanzees. These findings preliminarily support the idea that despite a wide geographical distribution throughout Africa, chimpanzees share some essential conventional behavioural patterns.     

Reproductive Parameters of Female<Emphasis Type="Italic">Pan paniscus</Emphasis> and <Emphasis Type="Italic">P. troglodytes</Emphasis>: Quality versus Quantity

We investigated intra- and interspecific differences in life history and reproductive parameters in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes). We compare the parameters of wild and captive females in order to shed light on the influence of habitat or specific differences or both on reproduction. We present new and additional information on reproductive parameters from captive bonobos and chimpanzees. Captive chimpanzees birth more live offspring and have a shorter interbirth interval, but experience higher infant mortality than captive bonobos. Although captive bonobo females tend to start reproduction at a younger age than chimpanzees, this is effectively only so for wild-born females of both species. Ultimately both species reach the same rate of production of offspring surviving to 5 yr. These results contrast with data from the wild. Wild bonobos tend to have higher reproductive success, a higher fertility rate and a shorter interbirth interval than wild chimpanzees. Reproduction is similar for wild and captive bonobos, which suggests that they are producing at their maximum under both conditions. Overall captive chimpanzees perform better than their wild conspecifics, probably because of lower feeding competition. Infant survival is the only specific difference not affected by captivity. Bonobo infants survive better, which suggests that chimpanzee infants are more at risk. We argue that the interspecific variation in reproductive parameters in captivity is related to the different influence of captivity on reproduction and different pressures of external sources of infant and juvenile mortality.     

On the possible adaptive value of coprophagy in free-ranging chimpanzees

Coprophagy occurred during major periods of feeding on fruits of Dialium spp. (Caesalpiniaceae) in a group of orphaned chimpanzees released in Conkouati Douli National Park, Republic of Congo. Since stress, boredom or food scarcity could not explain coprophagy according to our daily behavioral and veterinary control observations, we suggest that Dialium seeds were the item of interest in the feces. Two types of Dialium seeds were commonly found in the feces after chimpanzees swallowed the mesocarp and whole seeds together. These seeds were either whole and hard or whole/broken and soft imbibed. A mechanical and/or chemical effect of the gut passage may enable the chimpanzees to chew and ingest the seeds, thus providing nutritional intake.     

Physiological activities and the active constituents of potentially medicinal plants used by wild chimpanzees of the Mahale mountains,Tanzania

Potential medicinal plants for wild chimpanzees have been studied in order to discover their physiologically active compounds. Tests of the physiological activity of 3 plant species—Vernonia amygdalina, Aspilia mossambicensis, andFicus exasperata—indicate that they contain a variety of active compounds. From one species,V. amygdalina, an antitumor agent and 2 possible antitumor promoters are identified. Furthermore, steroid glucosides were isolated as the bitter substances. These structurally new compounds are expected to exhibit a number of significant physiological activities. The chemical investigation of possible medicinal plants used by chimpanzees should be helpful in recovering naturally occurring compounds of medicinal significance for human use.