Longitudinal changes in the targets of chimpanzee (Pan troglodytes) hunts at Mahale Mountains National Park: how and why did they begin to intensively hunt red colobus (Piliocolobus rufomitratus) in the 1980s?

The hunting activities of chimpanzees (Pan troglodytes) at Mahale Mountains National Park exhibited a significant change over a 46-year observation period, shifting from sporadic hunting for small ungulates and primates through seizure or chasing, to a specialized hunting habit for red colobus (Piliocolobus rufomitratus). By the early 1980s, a new hunting habit had developed through the following stages: (1) occasional single kills by mature/adolescent males, (2) multiple kills by mature/adolescent males, (3) participation of mature females in both multiple and single kills, and (4) increased frequencies of multiple kills and female participation in killing. Throughout the observation period, red colobus was the most frequently hunted species, accounting for 71.0% of all hunts, 68.1% of all kills, and 70.3% of the consumed prey. Adult/adolescent female chimpanzees accounted for 23.1% of all kills and 20.4% of red colobus kills, both of which are higher than proportions reported in other areas. There are several possible explanations for these changes: (1) improved observation conditions, including familiarization of the chimpanzees with human observers; (2) environmental changes such as forest and faunal recovery, particularly an increase in the red colobus population; (3) change(s) in observation methods; (4) a new invention or (5) reinvention by the Mimikire (M) group chimpanzees; and (6) the change in intergroup relationships between the M and Kajabala (K) groups.

Wild chimpanzees deprived a leopard of its kill: Implications for the origin of hominin confrontational scavenging.

This study reports the first observed case of wild chimpanzees (Pan troglodytes) obtaining animal prey freshly killed by a sympatric leopard (Panthera pardus) and scavenging it with the leopard still nearby. This observation has important implications for the emergence of confrontational scavenging, which may have played a significant role in human evolution. Many scholars agree that eating meat became important during human evolution, and hominins first obtained meat by scavenging. However, it is debatable whether scavenging behavior was "passive" or "confrontational (power)." The latter is more dangerous, as it requires facing the original predator, and it is thus considered to have been important for the evolution of several human traits, including cooperation and language. Chimpanzees do scavenge meat, although rarely, but no previous evidence of confrontational scavenging has hitherto emerged. Thus, it was assumed that they are averse to confrontation with even leopard-sized predators. However, in the observed case the chimpanzees frequently emitted waa barks, which indicated that they were aware of the leopard's presence but they nevertheless continued to eat the scavenged meat. In addition, we compiled and reviewed 49 cases of chimpanzee encounters with animal carcasses in the Mahale Mountains of Tanzania in 1980-2017. Chimpanzees scavenged meat in 36.7% of these cases, and tended to eat the meat when it was fresh or if the animal species was usually hunted by chimpanzees. However, no evidence indicated that carcasses were avoided when leopard involvement was likely. These results suggest that chimpanzee-sized hominins could potentially confront and deprive leopard-size carnivores of meat.

Brief communication: Orphaned Male Chimpanzees die young even after weaning.

If a social-living animal has a long life span, permitting different generations to co-exist within a social group, as is the case in many primate species, it can be beneficial for a parent to continue to support its weaned offspring to increase the latter's survival and/or reproductive success. Chimpanzees have an even longer period of dependence on their mothers' milk than do humans, and consequently, offspring younger than 4.5-5 years old cannot survive if the mother dies. Most direct maternal investments, such as maternal transportation of infants and sharing of night shelters (beds or nests), end with nutritional weaning. Thus, it had been assumed that a mother's death was no longer critical to the survival of weaned offspring, in contrast to human children, who continue to depend on parental care long after weaning. However, in theory at least, maternal investment in a chimpanzee son after weaning could be beneficial because in chimpanzees' male-philopatric society, mother and son co-exist for a long time after the offspring's weaning. Using long-term demographic data for a wild chimpanzee population in the Mahale Mountains, Tanzania, we show the first empirical evidence that orphaned chimpanzee sons die younger than expected even if they lose their mothers after weaning. This suggests that long-lasting, but indirect, maternal investment in sons continues several years after weaning and is vital to the survival of the sons. The maternal influence on males in the male-philopatric societies of hominids may be greater than previously believed.

Ranging behavior of Mahale chimpanzees: a 16 year study.

We have analyzed the ranging patterns of the Mimikire group (M group) of chimpanzees in the Mahale Mountains National Park, Tanzania. During 16 years, the chimpanzees moved over a total area of 25.2 or 27.4 km(2), as estimated by the grid-cell or minimum convex polygon (MCP) methods, respectively. Annually, the M group used an average of 18.4 km(2), or approximately 70 %, of the total home-range area. The chimpanzees had used 80 % of their total home range after 5 years and 95 % after 11 years. M group chimpanzees were observed more than half of the time in areas that composed only 15 % of their total home range. Thus, they typically moved over limited areas, visiting other parts of their range only occasionally. On average, the chimpanzees used 7.6 km(2) (in MCP) per month. Mean monthly range size was smallest at the end of the rainy season and largest at the end of the dry season, but there was much variability from year to year. The chimpanzees used many of the same areas every year when Saba comorensis fruits were abundant between August and January. In contrast, the chimpanzees used several different areas of their range in June. Here range overlap between years was relatively small. Over the 16 years of the study we found that the M group reduced their use of the northern part of their range and increased their frequency of visits to the eastern mountainous side of their home range. Changes in home-range size correlated positively with the number of adult females but not with the number of adult males. This finding does not support a prediction of the male-defended territory model proposed for some East African chimpanzee unit-groups.

Prevalence of muzzle-rubbing and hand-rubbing behavior in wild chimpanzees in Mahale Mountains National Park, Tanzania.

In 1998, four chimpanzees in the Mahale Mountains National Park, Tanzania, were observed wiping their mouths with non-detached leaves or stalks of grass, or rubbing their mouths with a tree trunk or branch, especially while eating lemons. The number of mouth-wiping/rubbing individuals increased to 18 in 1999. By 2005, 29 chimpanzees were documented wiping/rubbing their muzzles in this way. Although it is difficult to determine whether the chimpanzees acquired this behavior as a result of trial and error or social learning, the fact that chimpanzees at other sites perform this behavior with detached leaves or leafy twigs much more often than with intact items suggests the possibility that cleaning with intact plant parts at Mahale spread via social learning.

Sniffing behaviors in Mahale chimpanzees.

Although it is difficult for observers to determine how non-human primates use olfaction in a natural environment, sniffing is one clue. In this study, the sniffing behaviors of wild chimpanzees were divided into six categories, and sex differences were found in most categories. Males sniffed more frequently than females in sexual and social situations, while females sniffed more often during feeding and self-checking. Chimpanzees sniffed more frequently during the dry season than during the wet season, presumably due to the low humidity. This suggests that the environment affects olfactory use by chimpanzees and that chimpanzees easily gather new information from the ground via sniffing.

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.

Demography, female life history, and reproductive profiles among the chimpanzees of Mahale.

Demography provides critical data to increase our understanding of the evolution, ecology, and conservation of primate populations. The chimpanzees of the Mahale Mountains National Park, Tanzania, have been studied for more than 34 yr on the basis of individual identification and standardized attendance records. From this long-term study, we derived the following demographic data: The major cause of death was disease (48%), followed by senescence (24%) and within-species aggression (16%). Fifty percent of Mahale chimpanzees died before weaning. The median ages of female life history variables were: first maximal swelling, 10.0 yr (n = 5); emigration, 11.0 yr (n = 11); and first birth, 13.1 yr (n = 5). The median period of adolescent infertility was 2.8 yr (n = 4) when calculated from the age at immigration to that at first birth. Female fecundity was highest between 20 and 35 yr of age, with an annual birth rate of 0.2. Twenty-six females that were observed from a young age (10-13 yr) to death at various ages (15-40 yr) gave birth to an average of 3.9 and weaned an average of 1.4 offspring. Twenty-five females that were observed from middle age (18-33 yr) to death in older age (31-48) gave birth to an average of 2.7 and weaned an average of 2.0 offspring. The post-reproductive lifespan for female chimpanzees was defined as the number of years that passed from the year when the last offspring was born to the year when the female died, minus 5. Twenty-five percent of old females had a post-reproductive lifespan. The interbirth interval after the birth of a son (x = 72 mo) tended to be longer than that after the birth of a daughter (x = 66 mo). The extent of female transfer, which is a rule in chimpanzees, is influenced by the size and composition of the unit group and size of the overall local community.