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.     

Geographic variation in the calls of wild chimpanzees: A reassessment

Male chimpanzees produce a species‐typical call, the pant hoot, to communicate to conspecifics over long‐distances. Calls given by males from the well‐known Gombe and Mahale populations typically consist of four different phases: an introduction, build‐up, climax, and let‐down. Recent observations suggest that chimpanzees living in the Kibale National Park, Uganda, consistently give calls that lack a build‐up and are thus qualitatively distinguishable acoustically from those made by other East African conspecifics. We analyzed additional recordings from Mahale and Kibale to re‐examine geographic variation in chimpanzee calls. Results indicate that males from both sites produce pant hoots containing all four parts of the call. Calls made by chimpanzees from the two populations, however, differ in quantitative acoustic measures. Specifically, males at Kibale initiate their calls with significantly longer elements and build‐up over briefer periods at slower rates than individuals from Mahale. Kibale males also deliver acoustically less variable calls than chimpanzees at Mahale. Although climax elements do not differ between populations in any single acoustic feature, discriminant function analysis reveals that acoustic variables can be used in combination to assign calls to the correct population at rates higher than that expected by chance. Ecological factors related to differences in habitat acoustics, the sound environment of the local biota, and body size are likely to account for these observed macrogeographic variations in chimpanzee calls. Am. J. Primatol. 47:133–151, 1999. © 1999 Wiley‐Liss, Inc.     

One observed case of temporary adoption of an infant by unrelated nulliparous females among wild chimpanzees in the Mahale Mountains,Tanzania

A 2.4-year-old male chimpanzee in the Mahale Mountains, Tanzania became separated from his mother probably because of contagion of the flu-like sickness. He was adopted alternately by two unrelated immigrant females for at least six days before he was reunited with his mother. This episode is described and analyzed in relation to the babysitter relationships among female chimpanzees. The adaptive significance of the babysitting behavior of nulliparous females is discussed.     

Scavenging by chimpanzees at Ngogo and the relevance of chimpanzee scavenging to early hominin behavioral ecology

Chimpanzees regularly hunt a variety of prey species. However, they rarely scavenge, which distinguishes chimpanzee carnivory from that of some modern hunter-gatherers and, presumably, at least some Plio-Pleistocene hominins. I use observations made over an 11-year period to document all known opportunities for scavenging encountered by chimpanzees at Ngogo, Kibale National Park, Uganda, and describe all cases of scavenging. I also review data on scavenging from other chimpanzee research sites. Chimpanzees at Ngogo encountered scavenging opportunities only about once per 100 days and ate meat from scavenged carcasses only four times. Scavenging opportunities are also rare at other sites, even where leopards are present (Mahale, Ta?, Gombe), and scavenging of leopard kills is known only from Mahale. Feeding on prey that chimpanzees had hunted but then abandoned is the most common form of scavenging reported across study sites. For example, several individuals at Ngogo ate meat from a partially consumed red colobus carcass abandoned after a hunt the previous day. Such behavior probably was not common among Oldowan hominins. Ngogo data and those from other sites also show that chimpanzees sometimes eat meat from carcasses of prey that they did not see killed and that were not killed by chimpanzees, and that scavenging allows access to carcasses larger than those of any prey items. However, chimpanzees ignore relatively many opportunities to obtain meat from such carcasses. Scavenging may be rare because fresh carcasses are rare, because the risk of bacterial infections and zoonoses is high, and because chimpanzees may not recognize certain species as potential prey or certain size classes of prey species as food sources. Its minimal nutritional importance, along with the absence of technology to facilitate confrontational scavenging and rapid carcass processing, apparently distinguishes chimpanzee foraging strategies from those of at least some Oldowan hominins.     

The foraging adaptation of chimpanzees,and the recent behaviors of the provisioned apes in Gombe and Mahale National Parks,Tanzania

Current understandings of chimpanzee behavior and social organization are based largely on twenty years of studies of provisioned chimpanzees in Gombe and Mahale National Parks. These data indicate that chimpanzees are aggressive, dominance-seeking and fiercely territorial. Reports from a number of naturalistic (nonprovisioning, unobtrusive) field studies contrast sharply. All report open groups of nonaggressive, nonhierarchical chimpanzees, which repeatedly fission for foraging and reunite as a larger social group. Because of the authority accorded the Mahale and Gombe reports, the naturalistic studies are often discounted. In this paper foraging theory is used to show that the recent behaviors of the artificially fed apes are maladaptive, while those of the nonprovisioned, wild chimpanzees are excellent foraging strategy. It is suggested, but not concluded, that the recent behaviors of the Gombe and Mahale chimpanzees may be frustration-induced responses to feeding methods which introduce a blockage between the apes and the desirable bait foods.     

Behavioral seasonality in Mahale chimpanzees

To analyze how the chimpanzees in the Mahale Mountains National Park, Tanzania, change their grouping pattern, activity budget, travel speed, and travel distance within an annual cycle, I divided 1-year data into four periods. The Mahale chimpanzees have the behavioral flexibility to adapt to various climates and exhibited at least three behavioral seasons. In the early wet season, chimpanzees formed a few, large parties, and spent much time feeding on insects and animal meat. In the early and late dry seasons, chimpanzees maintained party sizes as large as in the early wet season, and traveled distances similar to the early wet season, but spent the most time feeding and traveling within the year. By contrast, in the late wet season chimpanzee parties broke up into more numerous, small groups, and traveled slowly over shorter distances. Although time spent feeding and traveling were the same as that in the early wet season, time spent feeding on terrestrial herbaceous vegetation (THV) was the highest in the year. The results suggest that chimpanzees travel longer, faster, and farther in seasons when they form large parties.     

Responses of wild chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>) toward seismic aftershocks in the Mahale Mountains National Park,Tanzania

This is the first report documenting the responses of wild chimpanzees (Pan troglodytes schweinfurthii) to seismic activities. During our long-term fieldwork in Mahale Mountains National Park, Tanzania, a high-intensity earthquake with a Richter magnitude of 6.8 occurred at 15:19 hours local time on 5 December 2005. During the main tremor, the chimpanzees displayed the “wraa” call, “scream,” and “pant bark” or “bark” vocalizations. Many mild aftershocks followed the main tremor, and the wild chimpanzees displayed a variety of responses to these. In several cases, they climbed trees or stopped activities such as grooming, moving, and feeding. These responses are similar to those previously reported in nonhuman primates. During the observations, a unique behavior, one never reported before was exhibited by a female chimpanzee. She placed her right palm on the ground giving the impression she was inspecting the trembling of the ground.     

A method of identifying chimpanzee hairs in lion feces

Hairs were detected in the feces of wild lions collected at the Mahale Mountains National Park, Tanzania. The materials were examined for their scale pattern, medulla arrangement, and ultrastructure of longitudinal sections. The observed features coincided with those of the chimpanzee, so that the hairs were judged to be from chimpanzees. The present results suggest that black hairs in lion feces can be identified as belonging to chimpanzees based on observations by scanning electron microscopy.     

Tool-assisted predation on a squirrel by a female chimpanzee in the Mahale Mountains,Tanzania

A 12-year-old female chimpanzee (Pan troglodytes schweinfurthii) of M-Group in the Mahale Mountains National Park, Tanzania was seen to rouse, capture, and eat a squirrel hiding in the narrow hole of a tree. The kill was aided by the use of a sturdy tool modified from a branch of the same tree. This appears to be the first reported case for chimpanzees, or any other nonhuman primate, of tool-use that directly led to the capture of a mammalian prey species. This behavior is discussed in relation to possible factors contributing to the occurrence of tool-use in small mammal predation especially by females to exploit a low competition meat source.     

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”.     

Role of Nonbehavioral Factors in Adjusting Long Bone Diaphyseal Structure in Free-ranging <Emphasis Type="Italic">Pan troglodytes</Emphasis>

Limb bones deform during locomotion and can resist the deformations by adjusting their shapes. For example, a tubular-shaped diaphysis best resists variably-oriented deformations. As behavioral profiles change during adulthood, patterns of bone deformation may exhibit age trends. Habitat characteristics, e.g., annual rainfall, tree density, and elevation changes, may influence bone deformations by eliciting individual components of behavioral repertoires and suppressing others, or by influencing movements during particular components. Habituated chimpanzee communities provide a unique opportunity to examine these factors because of the availability of morphological data and behavioral observations from known-age individuals inhabiting natural habitats. We evaluated adult femora and humeri of 18 female and 10 male free-ranging chimpanzees (Pan troglodytes) from communities in Gombe (Tanzania), Mahale Mountains (Tanzania), and Taï Forest (Côte d’Ivoire) National Parks. We compare cross sections at several locations (35%, 50%, 65% diaphyseal lengths). Community comparisons highlight different diaphyseal shapes of Taï females relative to Mahale and Gombe females, particularly in humeral diaphyses. Age trends in diaphyseal shapes are consistent with reduced activity levels in general, not only reduced arboreal activity. Age-related bone loss is apparent among community females, but is less striking among males. Community trends in diaphyseal shape are qualitatively consistent with ranked annual rainfall at localities, tree density, and elevation change or ruggedness of terrain. Habitat characteristics may contribute to variation in diaphyseal shape among chimpanzee communities, much like among modern human groups, but verification awaits further rigorous experimental and comparative analyses.     

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.     

Association partners of young chimpanzees in the Mahale Mountains National Park,Tanzania

Association partners of young chimpanzees at the Mahale Mountains National Park were analyzed. Juvenile and adolescent chimpanzees associated frequently with their mothers, although mother-offspring association decreased as the offspring grew up. Males tended to leave their mothers and associate with adult males, while females remained frequently associating with their mothers in early adolescence. In late adolescence and young adulthood, males usually associated with adult males and cycling adult females. Females may transfer into neighboring unit-groups in this stage. Although an immigrant female tended to be alone when her estrous cycle stopped, she associated with many individuals, in particular with adult males, when she resumed cycling. Some orphans were observed to associate frequently with particular adults. The findings were discussed in relation to the unique characteristics of chimpanzee social system.     

The leaf-clipping display: A newly-discovered expressive gesture in wild chimpanzees

Chimpanzees of the Mahale Mountains, Tanzania were discovered to show an expressive gesture using leaves, termed as “leaf-clipping display”. This behavioural signal is directed by an adult male to an estrous female as a possessive behaviour, or by an adolescent male as a courtship display, or by an estrous female to an adolescent male also as a solicitation of copulation. The signal also is used toward human observers as a signal of food-demanding. This behaviour pattern might originate in a displacement tool-making behaviour in conflict situations. The leaf-clipping display has not been observed in any other chimpanzee populations studied, and may probably be one example of the tradition drift in wild chimpanzees.     

Abundance and dispersion of some chimpanzee-dispersed fruiting plants at Mahale, Tanzania

Abundance and dispersion of five fruiting plant species the seeds of which are commonly dispersed by chimpanzees were investigated at Mahale Mountains National Park, Tanzania from August 2000 to February 2001. All plant individuals encountered within twenty‐three 50 × 50 m plots were counted and allocated to demographic stages. Plots were allocated proportionally to the main vegetation types in the chimpanzee range. Density and the demographic stages of individuals within 50 m radii of 10 adult conspecifics for four of the five sample species were also investigated. Overall, densities of individuals varied greatly by species and growth stages within the chimpanzee ranging space. Dispersion of the five plant species tested, irrespective of growth stages, showed a clumped pattern. However, Saba comorensis was the most evenly dispersed, irrespective of growth stage. Densities of juveniles and adults generally declined away from adult conspecifics, except for Syzygium guineense. These results are discussed in view of chimpanzees as potential seed dispersers. The hyper‐dispersion of S. comorensis appears to mirror the profound intensive and extensive dispersal of its seeds by chimpanzees, which are the main eaters of this fruit in the study area.     

Mortality rates among wild chimpanzees

In order to compare evolved human and chimpanzees' life histories we present a synthetic life table for free-living chimpanzees, derived from data collected in five study populations (Gombe, Ta?, Kibale, Mahale, Bossou). The combined data from all populations represent 3711 chimpanzee years at risk and 278 deaths. Males show higher mortality than females and data suggest some inter-site variation in mortality. Despite this variation, however, wild chimpanzees generally have a life expectancy at birth of less than 15 years and mean adult lifespan (after sexual maturity) is only about 15 years. This is considerably lower survival than that reported for chimpanzees in zoos or captive breeding colonies, or that measured among modern human hunter-gatherers. The low mortality rate of human foragers relative to chimpanzees in the early adult years may partially explain why humans have evolved to senesce later than chimpanzees, and have a longer juvenile period.     

Genital Ureaplasmas in nonhuman primates.

Ureaplasmas were isolated from the genital tracts of four of 22 (18.4%) male chimpanzees and eight of 23 (34.8%) female chimpanzees. Twenty-nine female rhesus monkeys, 38 female baboons, one gibbon, and black ape and one Java monkey were shown to be free of genital Ureaplasmas. The rate of reproductive failure among the chimpanzees was high and it is suggested that Ureaplasma may be responsible in part. The chimpanzee may serve as a useful model for human Ureaplasma genital infections.     

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.     

Body weights of wild chimpanzees (Pan troglodytes schweinfurthii) of the Mahale Mountains National Park, Tanzania

Ten male and nine female habituated chimpanzees (Pan troglodytes schweinfurthii) in the Kasoje area of the Mahale Mountains National Park, Tanzania, were weighed intermittently between December 1973 and March 1980 by luring them up a rope hung on a spring balance: six adult males averaged 42.0 kg and eight adult females 35.2 kg. Seasonal change in body weight was recognized at least partially; body weights tended to decrease in the later part of the wet season presumably because of food shortage in the middle of the wet season. Comparison of body weight among three populations of the same subspecies suggests that adult female chimpanzees of Mahale appear to be heavier than those of the Gombe National Park, Tanzania, and that they seem to be similar to the forest-living counterparts of eastern Zaire. On the other hand, body weights of adult male chimpanzees from the three populations do not show significant differences. Perhaps feeding competition among adult females in a small, isolated habitat is more severe than that among adult males, which may result in the body weight reduction among adult female chimpanzees at Gombe.     

Tool use and tool making in wild chimpanzees

Reported incidences of tool use and tool making for three wild chimpanzee populations increase from Mahale (12 and 3 types of use and making, respectively), Gombe (16 and 3) to Ta? (19 and 6). Sticks are commonly used and prepared at all three sites. However, Ta? chimpanzees seem to perform more modifications on the material before using it. They are also the only chimpanzees seen to pound objects with tools and to combine two different tool uses to get access to one food item. Tool making is the rule for abundant material (grass, twigs), but appears to be rarer for scarce, hard material (clubs, stones). Factors involved in the acquisition and the benefit of tool use are discussed along with factors affecting the frequency and complexity of tool making in chimpanzees.