Kenji Kawanaka

Factors Affecting Party Size in Chimpanzees of the Mahale Mountains

We studied factors affecting party size and composition of wild chimpanzees at Mahale (M group) over an 11-month period. Parties with 1–5 individuals were most frequent (37.8%; 153/405 parties); they included 94.7% of all male parties (n = 76) and 81.3% of all female parties (n = 75). The median of monthly values was the standard for analysis. We divided the year into four periods based on the median size of monthly bisexual parties (30.9 individuals; includes both males and females): monthly bisexual party sizes were larger in May–June (period II) and October–January (period IV) and smaller in February–April (period I) and July–September (period III). Only bisexual parties changed in size with period. The number of fruit items (=species) eaten was fewer in periods II and IV when abundance per item appeared to be great. The sizes of bisexual parties, which included cycling females with maximal anogenital swelling, were larger, and their representation (%) in all bisexual parties was greater in periods III and IV. The numbers of both cycling females and cycling females with maximal anogenital swelling were also larger in periods III and IV. The percentage of cycling females with maximal anogenital swelling was greater in periods II and III. The results of this study and those of Nishida (1979) suggest that seasonal variation in party size of Mahale chimpanzees maintains a relatively consistent annual cycle. The factors assumed to affect party sizes are fruit availability and the presence of cycling females with maximal anogenital swelling.

The Early Hominid Plant-Food Niche: Insights From an Analysis of Plant Exploitation by Homo, Pan, and Papio in Eastern and Southern Africa [and Comments and Reply]

African plant-food genera exploited by Homo, Pan, and Papio have been catalogued and analyzed to provide an estimation of the size and composition of the fundamental plant-food niche of the early hominids. Results to date include recognition of more than 100 widely distributed African plant genera which are the best known candidates for plant-food exploitation by the Plio/Pleistocene hominids of eastern and southern Africa. An analysis of staples reveals that fruits would be the most common type of plant part contributing to the early hominid plant-food diet. Six plant genera (four providing edible fruits) are the first genera to be identified as members of the most probable early-hominid fundamental plant-food niche. Potential interspecies competition for plant-food staples has also been estimated. It is highly significant and must be considered in models predicting the realized niche of these primates and the early hominids.

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-groups range, i.e., the overlapping area of the two unit-groups 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.

Association, ranging, and the social unit in chimpanzees of the Mahale Mountains, Tanzania

In order to characterize the social unit in chimpanzees, about which several conflicting views have been proposed, the proximity matrix among 55 recognized chimpanzees and the range covered by each of them are examined, on the basis of data obtained at the Mahale Mountains during 12 months in 1978–1979. It is shown once again that chimpanzees have a bisexual social unit (unit-group). Two such unit-groups were detected in the study area. All animals belonged to one of the two unit-groups except a few cycling females (and a juvenile male accompanied by his cycling mother) which were seen to associate alternately with members of two neighboring unit-groups, covering a whole range of one or even two unit-groups. The problem of such females is discussed in relation to the spatial relationships between the two unit-groups. Reexamining the membership of a unit-group, it is demonstrated that a unit-group was most likely patrilineal. While nulliparous females transferred between unit-groups, parous females tended to remain in a unit-group where they first gave birth to infants and to have several offspring therein. This appeared significant for ensuring recruitment of members of the next generation to a patrilineal unit-group. Although some adult males left their natal unit-group, they never joined the other. Male departure from a unit-group seemed to be forced by the other males and to be the sociological equivalent of going into exile, which is unique in nonhuman primates.

Further studies on predation by chimpanzees of the Mahale Mountains

Chimpanzees of the M-group in the Mahale Mountains, Tanzania, were seen to commit predation 15 times between May 1978 and July 1979. The data appear to suggest that they come to perform predation frequently in recent years. However, for evaluating long-term changes in their predatory disposition, several factors must be considered, such as changes in research formation, in the degree of habituation of the chimpanzees and in the influence of humans on prey animals. These factors should be also considered when the findings of various populations are compared.It seems that an adult male can more easily control his desire for exclusive possession of meat than that of plant-food. Adult males have priority over females in obtaining meat, but all of the former cannot join the meat-eating cluster. Besides the dominance rank, dyadic relationships between them may regulate whether an adult male can join it. This suggests that all adult males in a unit-group may not be tied by an equal bond.The evolutionary implications of chimpanzee predation are discussed on the basis of the assumption that both chimpanzee predation and human hunting originated in some habit of a common ancestor.

Extraordinarily low bone mineral density in an old female chimpanzee (Pan troglodytes schweinfurthii ) from the Mahale Mountains National Park

We examined bone mineral density (BMD) of the femoral neck and lumbar vertebrae of four chimpanzee skeletons from Mahale Mountains National Park, Tanzania, and four captive ones, with a dual energy X-ray absorptiometer. The BMD of Wansombo, an old female chimpanzee from Mahale , was remarkably lower than the mean of the other six younger adult female chimpanzees and categorized as osteoporosis. Posture, locomotion, and trunk-sacral anatomy of chimpanzees may have prevented fractures in Wansombo, whose BMD was below human osteoporosis criteria.

Age differences in social interactions of young males in a chimpanzee unit-group at the Mahale Mountains National Park, Tanzania

The behavior and social interactions of young male chimpanzees were studied in relation to their age change. The data were obtained at the Mahale Mountains National Park, during a four-month period in 1986. Early adolescent males, becoming independent of their mothers, spend a long time near adults of both sexes. Late adolescent males are not tolerated by the senior males. Although such animals do not stop traveling together with their seniors, they are separated from the other members including the males of their own age class, and each of them lives a relatively lonely life. Where seniors are not nearby, they perform charging displays in front of estrous females. Young adult males tend to remain in the proximity of the alpha male, and can associate with their seniors without pant-grunting. Although some young adult males dominate over some senior males, increasingly performing charging displays, they do not appear to be permitted to associate intimately with their seniors; they are not yet considered to have attained social maturity. Prime and senior males are strongly bonded with one another, being able to associate intimately with those of their own or senior age classes including the alpha male. A young adult males rise in rank is not connected with joining the “adult male-cluster,” nor does a senior males decline necessarily means his dropping out from the cluster: the social position of male chimpanzees cannot be understood solely from their agonistic dominance rank. The alpha male plays a leading part in integrating the males of the unit-group. Young adult males and their seniors tend to associate most frequently with him, and all the males of the early adolescent or senior age classes pay attention to his movements.

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 Pearsons correlation coefficients and Kendalls 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 animals 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.

Age differences in spatial positioning of males in a chimpanzee unit-group at the mahale mountains National Park, Tanzania

Proximity partner choice by male chimpanzees of various age classes was analyzed in relation to their spatial positioning. Field work was carried out twice at the Mahale Mountains National Park, Tanzania. Proximity data were recorded at 3 and 10m from the focal animal. The data for the proximity between the focal male and other individuals allowed the males to be classified into two categories according to both criteria: early adolescence to young adult, and prime to old age. Between the males, the 3m proximity data permitted a classification into two categories as above, but those for 10m did not. These two spatial distances thus probably have different meanings for the males. The numbers of male proximity partners and proximity with the alpha male also allowed the males to be classified into two categories: early and late adolescence, and young adult to old age. Together, the above results support the classification of males into three age-graded categories: (1) early and late adolescence, (2) young adult, and (3) prime to old age. This does not arise because the males of each category form an age group. Prime or older males are most frequently in proximity, while their juniors consistently attempt to approach them. However, even prime or older males are not equally in proximity with one another. Their proximity partners change as time passes. Probably recognizing such changes, they form coalitions or are in rivalry. The sexual interest of adolescent males is probably a factor stimulating them to separate from their mothers, and to approach older males. Young adult males, even though physically mature, do not have equal proximity relations with older males. They are not yet sufficiently qualified to join the coalitions formed by their seniors.

A Case of Inter-unit-group Encounter in Chimpanzees of the Mahale Mountains

In the Mahale Mountains, Tanzania, a case of encounter between chimpanzees of different unit-groups was observed in June 1979, where two young adult males and two adult females of the M-group came across four adult females and a juvenile male of the K-group. While the participants did not explicitly show any agonistic behavior but greeted and groomed each other, they repeatedly defecated during the interactions. This clearly indicated that they were put under unusual strain, which could be considered to be brought on because they were cognizant of that the opponents were not the members of their own unit-group. It can be said, therefore, that in chimpanzees, even females identify themselves with a particular unit-group, and that a chimpanzee unit-group should be regarded as a bisexual unit.