Conflict and post-conflict behavior in a small group of chimpanzees

Chimpanzee research plays a central role in the discussions of conflict negotiation. Reconciliation, or the attraction and affiliation of former opponents following conflict, has been proposed as a central element of conflict negotiation in chimpanzees and various other taxa. In an attempt to expand the database of chimpanzee conflict resolution, conflict and post-conflict behavior were recorded for a small group of socially housed chimpanzees at the Chimpanzee and Human Communication Institute, at Central Washington University. Data were collected over six 6-week periods between 1997 and 2000, for a total of 840 hours of observation, resulting in a substantial post-conflict (PC) and matched control (MC) data set. The data demonstrate this group’s tendencies to maintain visual contact and closer proximity after conflicts. Dyadic corrected conciliatory tendencies ranged between 0 – 37.5% and averaged 17.25% across all dyads. Individual corrected conciliatory tendencies ranged between 5.8 and 32%. The results of this study combined with recent publications on captive and free-ranging chimpanzee post-conflict behavior suggest that variation in post-conflict behavior may be important to our understanding of chimpanzee conflict negotiation, and may also have implications for the design and management of captive chimpanzee enclosures and social groups, respectively.     

Environmental factors influencing the seasonality of estrus in chimpanzees

Although the energetics of the estrous cycle in primates is not well understood, evidence suggests that energy and nutrient acquisition influence ovulation and the timing of conception. Energy for estrus has to compete with energy allocated for cellular maintenance, thermoregulation, movement for food, and predation avoidance. While some chimpanzee (Pan troglodytes) populations do not have a seasonal birth period, evidence suggests that there is seasonality in the number of estrous females. Similarly, the onset of postpartum cycles has been reported to be seasonal. We used 33 months of data from the Taï National Park, Côte dIvoire, to examine how the number of estrous females in a given month was influenced by the abundance and distribution of food, diet, rainfall and temperature. In a second analysis, we examined if there was a seasonal effect on first estrous swellings in adolescent females and postpartum adult females. Results demonstrated that the number of females in estrous in a given month was positively related to food abundance and percent foraging time spent eating insects, and negatively related to mean rainfall in the two preceding months and the mean high temperature. The timing of first estrous swellings of postpartum females and prepartum young females was positively related to the food abundance, and negatively related to mean high temperature. These results showed that environmental conditions can seasonally limit the energetically demanding estrus cycle. The presence of estrous females increases gregariousness in chimpanzee communities, and this study identified environmental factors that affect estrus directly and hence social grouping indirectly.     

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.     

Hierarchy and social status in Budongo chimpanzees

The status hierarchy is fundamental in the lives of male chimpanzees. This study describes the dominance interactions and social status among adult male chimpanzees of the Sonso community in the Budongo Forest Reserve, Uganda, during the period that they were first studied (1994 and 1995). Social dominance is typically measured using the behaviour of either the subordinate or the dominant individual, but a relationship is dependent on the behaviour of both parties and this study explicitly used both subordinate and dominant behaviours to investigate the status hierarchy. Among adult males of the Sonso community, agonistic interactions occurred at a low rate and pant-grunts were rare, but males could be ranked into separate hierarchies of agonistic dominance and pant-grunting (labelled respect) using ratios of behaviour performed/behaviour received. These hierarchies were combined to form a single hierarchy of social status that divided the males among five distinct status levels. The highest status level was held by an alliance between two males who replaced the previous alpha male during the first part of the study. Neither male in this alliance partnership pant-grunted to the other, although the reason for cooperative behaviour was unclear. Although the nominally beta male was treated as such by other adult males, he achieved surprisingly little mating success. Budongo Forest chimpanzees do not warrant the sometimes-expressed view that they are non-aggressive and peaceable and the broad pattern of their status interactions matches with that seen in other chimpanzee populations.     

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.     

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.     

Nut cracking by wild chimpanzees in Sierra Leone,West Africa

I report an incidence of chimpanzee (Pan troglodytes verus) cracking ofDetarium senegalense (Caesalpiniaceae) nuts on Tiwai Island, Sierra Leone. Similar behavior has been reported from other West African locations, but not from Central or East Africa.     

Tool-set for termite-fishing and honey extraction by wild chimpanzees in the Lossi Forest, Congo

The use of perforating sticks and flexible stalks in combination for termite fishing and a complex tool-set of three components used sequentially (stout chiel, bodkin, and dip-stick) to penetrate melipone and ground-dwelling bee hives byPan troglodytes troglodytes are documented or, inferred from circumstantial evidence. Functionally, termite extraction tools were similar to other locations in west and central Africa, but the plants and the number of raw material species used were different. Tools varied in the degree of modification (fraying ends). Chimpanzees in the Lossi forest seem to be able to use the tools not in a stereotyped fashion, but in a flexible, insightful way. The extraction of Melipone honey using large pieces of wood as pounding tools has rarely been recorded elsewhere. The most impressive technological solution to the honey-getting problem by wild chimpanzees was shown by this study. This is the only known, use of a tool-set of three components in sequence to extract honey by wild chimpanzees.     

Observation time and sampling intervals for measuring behavior and interactions of chimpanzees in the wild

This study examined two problems in the measurement of chimpanzee behavior: (1) comparability among data sets varying in length of total observation time; and (2) the longest interval for scoring reliable numbers of sample points with instantaneous sampling (this required procedures for evaluating the chi-square statistics of the sampled data). During a 4.5-month field study conducted at the Mahale Mountains National Park, Tanzania, one adult male was observed as a focal animal for about 300 hr with continuous recording. His behavior was classified into five categories. Data sets varying in total time were prepared by extraction from the raw data. Comparability among the data sets was evaluated using Pearson's correlation coefficients and Kendall's coefficients of concordance calculated from two kinds of measures obtained from the raw and simulated data sets: (a) the percentages of time spent by the focal animal in each behavior category; and (b) those of the time spent by adult males in his proximity. The results revealed that observation time of 25 hr was the critical length for scoring the above measures reliably. Sample points for the focal animal's behavior categories and for adult males in his proximity were simulated with intervals of various lengths for data sets differing in total time. The longest interval was measured by comparing the simulated scores with confidence limits calculated for the number of sample points to be scored with the respective intervals. It was found that the interval for sampling should be set at 3 min or shorter, and that chi-square statistics calculated from the data sampled with such an interval should be evaluated after their modification into the values to be obtained from the data sampled with a 5-min interval. These results may not be directly applicable to studies dealing with other behavior categories, other age/sex classes of focal animals, etc. However, the above problems should be examined widely in studies attempting to measure animal behavior, and the methods employed in this study are applicable to such studies.     

Establishing the presence of a body temperature rhythm in chimpanzees (Pan troglodytes) using a tympanic membrane thermometer

Seven chimpanzees (Pan troglodytes) were trained to present their ears so that a tympanic nembrane thermometer could be inserted. Temperatures were collected from both ears of each subject every 3 hours for 72 consecutive hours. The presence of a body temperature rhythm, well documented in other mammals, was established. Each ear demonstrated its own rhythm, but the rhythms in both ears generally mirrored each other. Similarities in the temperature rhythms of cagemates were found. These data are the first evidence of a body temperature rhythm in chimpanzees, and they represent a non-invasive method of measuring the 24-hr rhythms in both human and non-human primates.     

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.     

Locomotor behavior and long bone morphology in individual free-ranging chimpanzees

We combine structural limb data and behavioral data for free-ranging chimpanzees from Ta? (Ivory Coast) and Mahale National Parks (Tanzania) to begin to consider the relationship between individual variation in locomotor activity and morphology. Femoral and humeral cross sections of ten individuals were acquired via computed tomography. Locomotor profiles of seven individuals were constructed from 3387 instantaneous time-point observations (87.4 hours). Within the limited number of suitable chimpanzees, individual variation in locomotor profiles displayed neither clear nor consistent trends with diaphyseal cross-sectional shapes. The percentages of specific locomotor modes did not relate well to diaphyseal shapes since neither infrequent nor frequent locomotor modes varied consistently with shapes. The percentage of arboreal locomotion, rather than estimated body mass, apparently had comparatively greater biological relevance to variation in diaphyseal shape. The mechanical consequences of locomotor modes on femoral and humeral diaphyseal shapes (e.g., orientation of bending strains) may overlap between naturalistic modes more than currently is recognized. Alternatively, diaphyseal shape may be unresponsive to mechanical demands of these specific locomotor modes. More data are needed in order to discern between these possibilities. Increasing the sample to include additional free-ranging chimpanzees, or primates in general, as well as devoting more attention to the mechanics of a greater variety of naturalistic locomotor modes would be fruitful to understanding the behavioral basis of diaphyseal shapes.     

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.     

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.     

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.     

Preliminary report on predatory behavior and meat sharing in tschego chimpanzees (Pan troglodytes troglodytes) in the Ndoki forest,northern Congo

Predatory behavior ofPan t. troglodytes in the Ndoki Forest was confirmed by both direct observation and fecal evidence. Eight out of 214 fecal samples (3.7%), collected during 16 months, contained vertebrate tissue. The prey species were a terrestrial bird, two monkey species including crowned guenon, a squirrel, and probably a pangolin. This rate suggested that predation in the Ndoki population can occur as frequently as in other populations. Chimpanzees were also directly observed to eat an infant crowned guenon, a hornbill, and a duiker. An adult female used a branch apparently in an attempt to drive out a hornbill from its nest hole, though no bird was observed to come out. Chimpanzees were attracted to meat, and were observed begging and sharing over the meat. Predatory behavior is common toPan andHomo, but not toGorilla, implying that the common ancestor of the former two genera acquired this behavior after separating from gorillas.     

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.     

Wide-ranging chimpanzees at Mt. Assirik,Senegal

A 4-year study of the ecology and ethology of wild chimpanzees (Pan troglodytes verus)was carried out in far western Africa. Contacts with chimpanzees and the locations of their nests were noted to determine which types of habitat were most used and to estimate the density of the population and the size of its home range. The results show that this community has one of the lowest densities and largest home ranges of all populations of chimpanzees studied so far. As such, it may provide a useful model for the reconstruction of hominid evolution in the Plio-Pleistocene.