Chimpanzees' (Pan troglodytes) choice of prey among termites (macrotermitinae) in western Tanzania

This study examines factors affecting chimpanzees' preying on termites, especially their choice of prey species. Regular surveys of a large sample of termite-mounds in the Mahale Mountains, Tanzania, confirmed that chimpanzees of B Group obtainedMacrotermes, which were relatively uncommon, by the use of fishing-tools, but ignored the much more abundantOdontotermes. This preference appeared to reflectMacrotermes' more extended swarming period, greater accessibility, larger size, and less noxious taste than the soldiers ofOdontotermes. Similar factors probably underly the chimpanzees' choice ofMacrotermes at Gombe, butPseudacanthotermes spiniger are also common there, and their absence from the diet is less easily explained. In contrast, the chimpanzees of K Group at Mahale prey mainly onP. spiniger;Macrotermes are apparently absent, andP. spiniger appear to be more abundant, accessible, and palatable than the alternativeOdontotermes.     

Potential hominin plant foods in northern Tanzania: semi-arid savannas versus savanna chimpanzee sites

Savanna chimpanzees are useful as referential models for early hominins, and here potential differences between chimpanzee and early hominin ecology is the focus. Whereas chimpanzees inhabit only a handful of modern African savannas, there is evidence that early hominins occupied relatively more open and arid savannas than those in which chimpanzees live. In order to help expand potential models of early hominin palaeoecology beyond savanna chimpanzee-like scenarios, and to provide a basis for future modeling and testing of actual hominin diets, this study compares the types of plant foods available in modern semi-arid savannas of northern Tanzania to plant foods at savanna chimpanzee sites. The semi-arid savannas are not occupied by modern chimpanzees, but are potentially similar to environments occupied by some early hominins. Compared to savanna chimpanzee habitats, the northern Tanzania semi-arid savanna has a lower density and fewer species of trees that produce fleshy fruits. Additionally, the most abundant potential hominin plant foods are seasonally available Acacia seeds/pods and flowers, grass seeds, and the underground parts of marsh plants, as evidenced by vegetation surveys and by studies of the diets of baboons that forage in similar areas. The information from this study should be useful for framing hypotheses about hominin diets for sites with palaeoenvironmental contexts similar to those of the northern Tanzania semi-arid savannas and for contextualising tests of actual hominin diets (e.g., those based on dental microwear or isotopes).     

Inter-menstrual intervals in captive bonobosPan paniscus

In this study we provide new data on the duration of the inter-menstrual intervals of six captive female bonobos (Pan paniscus). We found that the mean duration of the inter-menstrual interval was about 34 days. This lies close to the average value of 37 days that has been reported for common chimpanzees (Pan troglodytes).     

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.     

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.     

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.     

A comparison of buttress drumming by male chimpanzees from two populations

Wild chimpanzees (Pan troglodytes) produce low-frequency sounds by hitting the buttresses and/or trunks of trees. This buttress drumming occurs in discrete bouts that may be integrated into the phrase sequence of the chimpanzees long-distance vocalization, the pant hoot. The aim of this study was to investigate whether regional variation exists in the drumming behavior of male chimpanzees from Kibale National Park (Kanyawara community), Uganda, and Taï National Park, Ivory Coast. Recordings were made during a 6-month field season at Taï in 1990, and a 12-month field season at Kanyawara in 1996–1997. Acoustic analysis revealed the following: (1) Kanyawara males drummed significantly less frequently in conjunction with a pant hoot or hoot than did Taï males; (2) drumming bouts by Kanyawara males included significantly fewer beats, and were significantly shorter in duration, than those of Taï males; these differences disappeared when only those bouts produced in conjunction with a call were compared; (3) when Kanyawara chimpanzees did call and drum together, they tended to integrate drumming into the vocalization at a later point than did Taï males; and (4) individual differences in the temporal patterning of drumming bouts were not apparent for Kanyawara males, whereas a previous analysis revealed individual differences among Taï males.     

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.     

Prospects for Bonobo Insectivory: Lui Kotal, Democratic Republic of Congo

Chimpanzees (Pan troglodytes) are well-known to eat invertebrates, especially social insects, across Africa, but allopatric bonobos (P. paniscus) are not. Bonobo insectivory is sparsely documented and apparently sporadic. However, the availability to bonobos of social insect prey and raw materials with which to make tools to exploit them is unknown. Here, we test a set of hypotheses that relates to questions of presence, abundance, density, and distribution of taxa that Pan consume and of vegetation suitable for making extractive foraging tools. We worked at Lui Kotal, Democratic Republic of Congo, where unprovisioned bonobos live in intact forest, far from villages. We collected insect and fecal specimens, transected for prey and assessed raw materials, and monitored mounds of Macrotermes. All but 1 of the major taxa of relevant termites, ants, and (stinging) honey bees were present. The 3 main taxa of insects that chimpanzees elsewhere eat —Macrotermes (fungus-growing termites), Dorylus (Anomma; army or driver ants), and Apis (honey bees)— were abundant and widespread, and usually at densities exceeding those at well-known chimpanzee study-sites. Similarly, woody and nonwoody vegetation suitable for making fishing probes was common at mounds of Macrotermes. There is no obvious ecological reason why bonobos should not use elementary technology in extractive foraging, e.g., termite-fish, ant-fish, ant-dip, honey-dip, to obtain social insects.     

A case of offspring desertion by a female chimpanzee and the behavioral changes of the abandoned offspring

In the Mahale Mountains National Park, Tanzania, a chimpanzee mother moved to a neighboring unit-group leaving her 5-year-old son behind in the original group. The mother remained in the new group for ten months and then after conceiving, returned to the original group. The mother and son were reunited again but the son died of illness two months later. The mother gave birth to a baby in the original group but eventually transferred to the neighboring group after that baby disappeared. This is the first record of an immature chimpanzee abandoned by its emigrating mother, although a similar phenomenon has been reported for gorillas and Hanuman langurs. The abandoned offspring followed an adult male of the unit-group and the male was the individual who gave most of the care to the offspring. The offspring's time spent on grooming others increased, the time spent on being groomed decreased, and the rate of aggression toward him increased after his mother's desertion. This report will occasion to reconsider the previous notion of all-powerful mother-offspring bond in chimpanzees.     

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.     

What chimpanzees (Pan troglodytes) learn in a cooperative task

To examine the development of cooperation in a captive group of chimpanzees (Pan troglodytes), we designed an apparatus which required the simultaneous traction by two animals to get a reward. Two chimpanzees, an adult male and an infant female in a group of six, produced most of the successful responses (pulling on two handles simultaneously). Visual behavior was used to try to determine what chimpanzees learned about the cooperative task. Propositions were made to investigate what kind of learning could be attributed to chimpanzees and were confronted with results. Both subjects learned the link between the presence of fruits on the apparatus and the possibility of getting a fruit. They also learned the importance of the partner at the apparatus to make a successful response. Only the adult male learned to take into account the behavior of the partner at the apparatus before pulling a handle. From a methodological point of view, the glances made by the animals can constitute a useful behavioral indicator of what the subjects learned in a given social situation.     

Within-group cannibalism by adult male chimpanzees

In the Mahale Mountains National Park, Tanzania, a young adult male chimpanzee was observed to feed on a 3-month-old male infant of the same unit-group. Four other adult males and an adult female shared the carcass. The mother of the victim had immigrated from a neighboring unit-group four years previously. Circumstantial evidence strongly suggests that the first-observed cannibal male also killed the infant. The adult male and the mother of the victim had been familiar socially and sexually with each other since the female immigrated. Since the mother of the victim had usually been ranging in the peripheral part of the unit-group's range, i.e., the overlapping area of the two unit-group's ranges during pregnancy and soon after birth, the infanticidal male might have had reason to suspect the paternity of her infant. Four such cases of within-group cannibalism by adult males suggest that the female range and association pattern before and after parturition are key factors allowing an infant to survive. The possibility of male-biased infanticide is also discussed.     

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.     

Body size and proportions in chimpanzees,with special reference toPan troglodytes schweinfurthii from Gombe National Park,Tanzania

Body weight, cranial capacity, linear and joint area data from ten free-ranging adult chimpanzees from Gombe National Park, Tanzania with known life histories allow study of variation in a local population and comparison to other populations ofPan troglodytes and toPan paniscus. Because individuals in the Gombe population are small compared to other common chimpanzees, they provide a useful comparison toPan paniscus. Body weight and some linear dimensions overlap withPan paniscus. However, cranial capacity, tooth size, and body proportions of Gombe individuals lie within the range of otherPan troglodytes and are distinct fromPan paniscus.     

An apparatus and procedure for studying the choice component of foraging in a captive group of chimpanzees (Pan troglodytes)

An apparatus is described which was used to investigate the choice component of foraging in a captive group of chimpanzees maintained in a large, outdoor compound at the Yerkes Regional Primate Research Center Field Station. The utilization of more than one apparatus would allow the investigation of other ecological and psychological concepts in nonhuman primates housed under semi-natural conditions.     

Personality in Captive Chimpanzees: Use of a Novel Rating Procedure

We present a new method to assess the personalities of captive chimpanzees (Pan troglodytes). Using individually generated bipolar constructs, experienced observers rated the personalities of chimpanzees on a five-point scale. Each observer chose a subset of his/her own constructs and we performed a separate factor analysis on each set of ratings. The analyses indicate four factors: dominance, sociability, machiavellianism, and anxiety. We propose that this factor structure will be useful in the development of a single rating instrument with which to assess the personalities of captive chimpanzees.     

Chimpanzees (Pan troglodytes) use markers to monitor the movement of a hidden item

Four chimpanzees (Pan troglodytes) monitored the movement of hidden items in arrays of opaque cups. A chocolate candy was hidden in an array of four cups and temporarily presented paper markers indicated the location of the candy (which otherwise was not visible). These markers were either non-symbolic or symbolic (lexigram) stimuli that in other contexts acted as a label for the hidden candy, and the array was either rotated 180° after the marker was removed or the array remained in the same location. For three of four chimpanzees, performance was better than chance in all conditions and there was no effect of the type of marker. These experiments indicate that chimpanzees can track the movement of a hidden item in an array of identical cups even when they never see the item itself, but only see a temporarily presented marker for the location of that item. However, there was no benefit to the use of symbolic as opposed to non-symbolic stimuli in this performance.     

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.