Hunting Behavior of Chimpanzees at Ngogo,Kibale National Park,Uganda

Chimpanzees (Pan troglodytes) prey on a variety of vertebrates, mostly on red colobus (Procolobus spp.) where the two species are sympatric. Variation across population occurs in hunting frequency and success, in whether hunting is cooperative, i.e., payoffs to individual hunters increase with group size, and in the extent to which hunters coordinate their actions in space and time, and in the impact of hunting on red colobus populations. Also, hunting frequency varies over time within populations, for reasons that are unclear. We present new data on hunting by chimpanzees at Ngogo, Kibale National Park, Uganda, and combine them with earlier data (Mitani and Watts, 1999, Am. J. Phys. Anthropol. 109: 439–454) to examine hunting frequency and success, seasonality, and cooperation. The Ngogo community is the largest and has the most males of any known community. Chimpanzees there mostly hunt red colobus and are much more successful and make many more kills per hunt than at other sites; they kill 6–12% of the red colobus population annually. The number of kills and the offtake of meat per hunt increase with the number of hunters, but per capita meat intake is independent of hunting party size; this suggests that cheating occurs in large parties. Some behavioral cooperation occurs. Hunting success and estimated meat intake vary greatly among males, partly due to dominance rank effects. The high overall success rate leads to relatively high average per capita meat intake despite the large number of consumers. The frequency of hunts and of hunting patrols varies positively with the availability of ripe fruit; this is the first quantitative demonstration of a relationship between hunting frequency and the availability of other food, and implies that the chimpanzees hunt most when they can easily meet energy needs from other sources. We provide the first quantitative support for the argument that variation in canopy structure influences decisions to hunt red colobus because hunts are easier where the canopy is broken.     

Intracommunity Coalitionary Killing of an Adult Male Chimpanzee at Ngogo,Kibale National Park,Uganda

Intercommunity coalitionary killing of adult and adolescent males has been documented in two chimpanzee communities in the wild, and it was strongly suspected in a third. It may increase survivorship for the attackers, their mates, and their offspring by reducing the combined strength of hostile neighbors and/or by increasing territory size and food availability, and it may help the attackers to attract mates. Lethal coalitionary attacks by males on other male members of their own communities would not provide these benefits and are not expected, given the importance of cooperation among male community members in contests for dominance rank and in both defense and offense against neighboring males. Nevertheless, intracommunity coalitionary killings associated with struggles for alpha rank occur in the wild and in captivity, and observers have seen serious gang attacks on maturing adolescent and young adult males at Mahale and Budongo: the victim in the Budongo case was killed (Fawcett and Muhumuza, 2000). I describe a lethal attack on a young adult male by a large coalition of males from his community at Ngogo, Kibale National Park, Uganda. The Ngogo community is the largest known for chimpanzees and has an unusually large number of males. The attack was not related to a struggle for alpha rank: the victim was low-ranking and the community had a well-established alpha at the time. However, the victim had risen substantially in the male hierarchy over the past few years and might have appeared threatening to many higher-ranking males. Simultaneously, he associated relatively little with most other adult males, had relatively few grooming partners and was not well integrated into the male grooming network, and had no influential allies. The combination of these social factors with the unusual demographic circumstances – which presumably meant that mating competition was relatively high and the cost of losing one male relatively low – might have triggered the attack.     

Variation in diet and ranging of black and white colobus monkeys in Kibale National Park,Uganda

Recently, considerable intraspecific variation in the diets and ranging behavior of colobine monkeys has been described, although in most cases this has involved documenting variation between, not within, sites. Some African colobines, such as guerezas (Colobus guereza), are relatively abundant in disturbed habitats that are very heterogeneous, raising the intriguing possibility that even groups with overlapping home ranges may exhibit large behavioral differences. If such differences occur, it will be important to understand what temporal and spatial scales adequately portray a species’ or population’s diet and ranging behavior. This study documents within-site variation in the diet and ranging behavior of guerezas in the habitat types in which they are described to be most successful—forest edge and regenerating forest. We collected data on eight groups of guerezas with overlapping home ranges for 3–5 months each in Kibale National Park, Uganda. The guerezas were highly folivorous, with leaves constituting 78.5–94.0% of the groups’ diets. The percentage of mature leaves and fruit in the diet varied widely among and within groups. We show that differences among groups in the intensity with which they fed on specific tree species were not just related to phenology, but also to differences in the forest compositions of groups’ core areas. Range size estimates varied more than fivefold among groups and the minimum distance from groups’ core areas to eucalyptus forest (which all groups regularly fed in) was a better predictor of range size than was group size. These results reveal considerable variation in the diet and ranging behavior among groups with overlapping ranges and have implications for comparative studies, investigations of within- and between-group feeding competition, and the potential for populations to adapt to anthropogenic or natural environmental change.     

Coping with Forest Fragmentation: The Primates of Kibale National Park,Uganda

A goal of conservation biology is to determine which types of species are most susceptible to habitat disturbance and which types of disturbed habitats can support particular species. We studied 20 forest fragments outside of Kibale National Park, Uganda, to address this question. At each patch, we determined the presence of primate species, tree species composition, patch size, and distance to nearest patch. We collected demographic, behavioral, and dietary data for Abyssinian black-and-white colobus (Colobus guereza). Black-and-white colobus and red-tailed guenons (Cercopithecus ascanius) were in almost all fragments; Pennant's red colobus (Procolobus pennantii) and chimpanzees (Pan troglodytes) were in some fragments; and blue monkeys (Cercopithecus mitis) and gray-cheeked mangabeys (Lophocebus albigena) were absent from all fragments. No species characteristics—home range, body size, group size, or degree of frugivory—predicted the ability of species to live in patches. No characteristics of patches—area, distance to the nearest patch, distance to Kibale, or number of food trees present—predicted the presence of a particular species in a patch, but distance to Kibale may have influenced presence of red colobus. Black-and-white colobus group size was significantly smaller in the forest patches than in the continuous forest of Kibale. For a group of black-and-white colobus in one patch, food plant species and home range size were very different from those of a group within Kibale. However, their activity budget and plant parts eaten were quite similar to those of the Kibale group. The lack of strong predictive variables as well as differences between other studies of fragmentation and ours caution against making generalizations about primate responses to fragmentation.     

Grouping Patterns and Competition Among Female <Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis> at Ngogo,Kibale National Park,Uganda

In mammals, access to mates is probably the most important influence on male reproductive success, whereas foraging efficiency is probably the most important influence on female reproductive success Emlen and Oring (Science 197:215–223, 1977). Male chimpanzees (Pan troglodytes) are highly gregarious and form cooperative relationships with other males. In contrast, female social relationships vary within and between populations. Females in most East African populations, e.g., Gombe, Mahale, Kibale-Kanyawara, are less gregarious than males and spend most of their time alone or with only their dependent offspring. Researchers have attributed low female gregariousness to the high potential for feeding competition. I provide the first data on association patterns and agonistic interactions of female chimpanzees (Pan troglodytes schweinfurthii) from the unusually large Ngogo community, Kibale National Park, Uganda. Ngogo females were less gregarious than males, but spent a mean of 64% of their time in association with ≥1 other females and as much time in all-female parties as they did alone. Further, female dyads associated nonrandomly and they formed associative cliques. Association levels within cliques were similar to those among the relatively gregarious West African chimpanzee females at Taï (Pan troglodytes verus) and among bonobo (P. paniscus) females. Agonistic interfemale interactions were extremely rare, and monthly mean party size and the numbers of anestrous females per party do not correlate significantly with fruit availability. Thus, Ngogo females maintained relatively high levels of gregariousness, but avoided detrimental feeding competition by preferentially associating with a small subset of other community females.     

Infanticide and cannibalism by male chimpanzees at Ngogo, Kibale National Park, Uganda

Researchers have documented infanticide by adult males in four wild chimpanzee populations. Males in three of these have killed infants from outside of their own communities, but most infanticides, including one from Kanyawara, in Kibale National Park, Uganda, took place within communities. Here we report two new cases of infanticide by male chimpanzees at a second Kibale site, Ngogo, where the recently habituated chimpanzee community is the largest yet known. Both infanticides happended during boundary patrols, which occur at a high frequency there. Patrolling males attacked solitary females who were unable to defend their infants successfully. The victims were almost certainly not members of the Ngogo community. Males cannibalized both infants and completely consumed their carcasses. These observations show that infanticide by males is widespread in the Kibale population and that between-community infanticide also happens there. We discuss our observations in the context of the sexual selection hypothesis and other proposed explanations for infanticide by male chimpanzees. The observations support the arguments that infanticide has been an important selective force in chimpanzee social evolution and that females with dependent infants can be at great risk near range boundaries, but why male chimpanzees kill infants is still uncertain.     

Nutritional ecology of Preuss's red colobus monkey (Colobus badius preussi Rahm 1970) in Korup National Park, Cameroon

Studies on the nutritive ecology of red colobus monkeys (Colobus badius preussi) in Korup National Park, Cameroon, investigated food plants and the parts eaten as well as nutritive values of the various food items. Food plants were identified through direct observations and collected for laboratory determination of nutrient content. Food plants were analysed for crude protein (CP), crude fibre (CF), ether extract (EE), nitrogen‐free‐extractive (NFE) and total ash (A). A total of seventeen species belonging to nine different families were identified as food plants for red colobus. Food and feeding habits showed that the species were highly folivorous, tending to prefer the young leaves of food plants. Lecomtedoxa klaineana (27.0%) and Xylopia aethiopica (22.2%) of families Sapotaceae and Annonaceae, respectively, constituted about 50% of total food items consumed. Results of nutrient composition indicated individual nutrients varied significantly (P < 0.01) with food species.     

New cases of inter-community infanticide by male chimpanzees at Ngogo,Kibale National Park,Uganda

Infanticide by males has been recorded in four chimpanzee populations, including that in Kibale National Park, Uganda. Some infanticidal attacks occur during inter-community aggression. The sexual selection hypothesis does not easily explain these attacks because they may not directly increase male mating opportunities. However, females in the attackers’ community may benefit by expanding their foraging ranges and thereby improving their reproductive success; thus infanticide may increase male reproductive success indirectly. We report two new cases of infanticide by male chimpanzees at Ngogo, Kibale National Park. Like two previous cases, these occurred during a boundary patrol and were almost certainly between-community infanticides. The patrolling males attacked despite the proximity of males from the victims’ presumed community. This probably explains why, unlike the earlier cases, they did not completely cannibalize their victims. Such attacks seem to be relatively common at Ngogo and infanticide may be an important source of infant mortality in neighboring communities. Our observations cannot resolve questions about the sexual selection hypothesis. However, they are consistent with the range expansion hypothesis: the infanticides occurred during a period of frequent encounters between communities associated with a mast fruiting event, and Ngogo community members greatly increased their use of areas near the attack site during another mast fruiting event one year later. Our observations contribute to growing evidence that lethal intergroup aggression is a common characteristic of wild chimpanzee populations.     

Taphonomic analysis of skeletal remains from chimpanzee hunts at Ngogo, Kibale National Park, Uganda

This study provides a taphonomic analysis of the largest known sample of bone fragments collected from chimpanzee hunts. The entire sample consists of 455 bone fragments from 57 chimpanzee hunting episodes of 65 prey individuals at Ngogo, Kibale National Park, Uganda. It has low taxonomic diversity, consisting overwhelmingly of primates, especially red colobus monkeys. The age distribution of the prey remains is skewed towards pre-adults. Cranial bones are the dominant element, followed by long bones. Axial postcranial elements have low survivorship, with a complete absence of pre-caudal vertebrae. Bone is damaged in distinct ways, such as: destruction of long bone ends, typically with intact but chewed shafts; fragmentation and compression cracking of crania; and preservation of only the iliac blades of the innominates. Tooth marks are present but uncommon (4.4% of total NISP). These analyses enable us to: 1) describe and characterize consistent patterns of bone damage inflicted by chimpanzees across a much larger prey sample than has been previously studied; 2) make a preliminary comparison of the generalized chimpanzee taphonomic signature to that of leopard and eagle consumption of primates, as well as modern human consumption of small mammals; and 3) assess the utility of such samples for recognition of early hominin small mammal carnivory. We present a model that may be useful for detecting a pre-technological hominin carnivory and suggest some fossil locales at which close inspection of cercopithecoid remains for the above patterns might reveal traces of hominin hunting, though we caution that a pre-technological hominin hunted "assemblage" is not likely to be archaeologically visible.     

The influence of forest variation and possible effects of poaching on duiker abundance at Ngogo, Kibale National Park, Uganda

Duikers were censused at the Ngogo study area, Kibale National Park, Uganda, between July 2002 and August 2004. Censuses were conducted along three transects, of which, two (colonizing forests 1 and 2) were located in colonizing forests naturally replacing anthropogenic grasslands and one in old growth forest. Colonizing forest 1 was more prone to poaching than both colonizing forest 2 and the old growth forest that were closest to the research camp. Duikers that were actually sighted were identified to species, red or blue. However, on some occasions, duikers were detected by alarm calls and/or movements as they fled; these were simply recorded as duikers. Duiker abundance, regardless of species or mode of detection, was higher in colonizing forest 2 than colonizing forest 1 and the old growth forest. However, when the analysis was restricted only to duikers that were sighted, and hence identified to species, red duiker abundance was highest in colonizing forest 2 followed by the old growth forest and was lowest in colonizing forest 1; all these differences were significant. Blue duiker abundance was lowest in the old growth forest despite its proximity to the research camp; however, this was only significantly lower than in colonizing forest 2. Apart from colonizing forest 1, red duikers were significantly more abundant than blue duikers in the other two forest sections. This study suggests that forests colonizing anthropogenic grasslands may support more duikers than old growth forests; poaching in colonizing forest 1 has a severe impact on the duiker population and, red duikers are affected more severely by poaching than blue duikers.     

A preliminary analysis of wound care and other-regarding behavior in wild chimpanzees at Ngogo,Kibale National Park,Uganda

Primates - Caring for others is a key feature of human behavior. Mothers, fathers, siblings, grandparents, and other group members provide care in the form of provisioning, protection, and first...     

Fruit and seed ecology of wild robusta coffee (Coffea canephora Froehner) in Kibale National Park, Uganda

Coffee fruit production, frugivorous activities and seed longevity were investigated during two seasons. The average densities of fruiting coffee trees during the first sampling period ranged from four to seven per 25 m² and during the second sampling period, four fruiting trees per 25 m² were estimated. Fruiting coffee trees were significantly less than the non-fruiting trees ( P < 0.05, t -test). Periods of maximum coffee fruit fall coincided with those of lowest rainfall. Coffee fruit yield per tree from ground collections was 7–40 m⁻² in 1992 and 16–38 m⁻² in 1993/1994. Split coffee berries ranged between 71.7% and 83.6% of all fruits collected from the forest floor in the 1993/94 sampling period. Coffee seeds collected from the forest floor were mostly undamaged. Black and white colobus and redtail monkeys were observed to feed on ripe coffee fruits but did not crush the seeds. Coffee seed viability declined rapidly during storage. Seeds left on the forest floor survived longest, those stored under laboratory conditions lost viability fastest, and those in cold storage showed intermediate longevity.     

Spatial distribution of primates in a mosaic of colonizing and old growth forest at Ngogo, Kibale National Park, Uganda

Primate censuses were conducted in a mosaic of colonizing (two locations) and old-growth forests using line transect methods at the Ngogo study site, Kibale National Park, Uganda. Black and white colobus monkeys (Colobus guereza) were encountered more frequently in the colonizing forests than in the old growth forest, while chimpanzees (Pan troglodytes) were encountered more frequently in the old growth forest than in colonizing forests. Although not significant, results suggest that blue monkeys (Cercopithecus mitis) frequented colonizing forests more often than old growth forest. The encounter rates of mangabey (Lophocebus albigena), and redtail (Cercopithecus ascanius) groups were ambiguous with their density being higher in some colonizing forests but not others as compared to old-growth forest. No significant differences were detected for baboons (Papio anubis), Lhoests (Cercopithecus lhoesti), and red colobus monkeys (Piliocolobus tephroscales). The conversion of forests to farmland is one of the major problems encountered in primate conservation. This study shows that secondary forests replacing anthropogenic grasslands have the potential of supporting some primate species such as black and white colobus, redtail monkeys, and possibly blue monkeys. Therefore, such areas should not be given up but should be conserved for the benefit of primates that can survive in secondary forests; as the forests mature further, primate species that are adapted to old growth forest will colonize the area provided there is a nearby source.     

Grooming Between Male Chimpanzees at Ngogo,Kibale National Park. II. Influence of Male Rank and Possible Competition for Partners

Allogrooming contributes to the development and maintenance of social relationships, including those that involve alliances, in many primate species. Variation in relatedness, dominance rank, and other factors can produce variation in the value of others as grooming partners. Several models have been developed to account for variation in the distribution of grooming in relation to dominance ranks. These start from the premise that individuals are attracted to high-ranking partners, but time limits, direct competition, and prior grooming engagement between high-ranking individuals can constrain access to them. Sambrook et al. (1995) formalized some of these models and showed the importance of taking group size variation into account when assessing them. Chimpanzees form multimale communities in which males are the philopatric sex. Males commonly associate and groom with each other; they also form dominance hierarchies and form alliances that influence dominance ranks and mating success. Both male rank and the rank distance between partners are significantly correlated with the distribution of grooming between males in an extremely large chimpanzee community at Ngogo, Kibale National Park, Uganda, that has more males than any other known community. High-ranking males had more grooming partners than mid- or low-ranking males. Grooming predominantly went up the dominance hierarchy, but was also concentrated among males that were close in rank. Rank and rank distance apparently both affected grooming independently of reciprocity in grooming and independently of the frequency with which males associated in temporary parties. However, the data do not clearly indicate how constraints on access to partners might have operated. Published data from a smaller chimpanzee community at Mahale show no rank or rank distance effect on male grooming. These results and earlier, conflicting findings on the association between dominance rank and grooming in male chimpanzees indicate that variation in group size, i.e., the number of males per community, probably influences the strength of any such effects, as happens for grooming between females in several cercopithecine species. Data on coalitions at Ngogo support the argument that high-ranking males are valuable social partners, and similarity in strategies of alliance formation may influence the distribution of grooming.     

Species,age and sex differences in type and frequencies of injuries and impairments among four arboreal primate species in Kibale National Park,Uganda

Animals in the wild often have physical impairments that can affect their fitness. The aim of this study was to compare injuries and impairments of four different primate species (black-and-white colobus, red colobus, red-tailed monkeys, and grey-cheeked mangabeys) living in Kibale National Park (Uganda), and estimate the proportion of injured and impaired individuals among the sexes and age classes. The species differed in the proportion of individuals with injuries and impairments, with 16.7% in black-and-white colobus, 23.1% in red colobus, 16.2% in red-tailed monkey and 30.2% in grey-cheeked mangabeys. Species also differed in the types and location on the body of injuries. Adult animals had more injuries than immatures. Males had more injuries than females, in all but red-tailed monkeys. The results are discussed in relation to the literature on aggressive behavior, locomotion, and predation in these species.     

Effects of male group size,parity, and cycle stage on female chimpanzee copulation rates at Ngogo,Kibale National Park,Uganda

Chimpanzees have complex and variable mating strategies, but most copulations occur when females with full sexual swellings are in parties with multiple males and mate with most or all of those males. Daily copulation rates for fully swollen females vary at different times of a female’s cycle, among females, and across communities and populations. Variation in female age, parity, and cycle stage underlie some of this variation, but possible demographic effects on copulation rates have not been systematically investigated. Demographic variation can affect many aspects of behavior and ecology, including the frequency and success of different mating tactics. Analysis of data from the unusually large chimpanzee community at Ngogo produces two results that are consistent with the hypothesis that demographic variation affects female copulation rates. Copulation rates were high compared with those reported from other research sites, where females had fewer potential mates available. Daily copulation rates of fully swollen females were also positively related to the number of males with whom they associated. Ngogo data also re-confirm results from other studies, of both wild and captive populations, showing that female copulation rates increase during periovulatory periods. This is consistent with the hypothesis that sexual swellings and extended receptivity and proceptivity help to protect females against infanticide by helping to ensure they mate with all potential sires. As at some other sites, parous females at Ngogo copulated at higher rates than nulliparous females. Possible effects of demography on sexual behavior should be considered in assessments of differences between chimpanzees and bonobos and of variation across chimpanzee populations.     

Parallel lasers and digital photography to estimate limb size of chimpanzees (Pan troglodytes) at Ngogo,Kibale National Park,Uganda

Primates - How animals grow and when they stop growing are key variables for understanding life history evolution. Although theoretically straightforward, it is logistically difficult to take body...     

Grooming Between Male Chimpanzees at Ngogo,Kibale National Park. I. Partner Number and Diversity and Grooming Reciprocity

Allogrooming serves many social functions in primates. Grooming can help individuals to service social relationships generally, sometimes reciprocally, and may be particularly important in the development and maintenance of alliances. However, time constraints limit the number of partners with whom one individual can groom enough to maintain cooperative relationships. As a result, the size of its grooming network may reach an asymptote as the size of its group increases, and it may distribute its grooming less equally among potential partners. Chimpanzees live in multimale, fission-fusion communities; males are philopatric, and commonly associate and groom with each other. Males form within-community alliances that influence dominance rank and access to mates, and allies groom with each other regularly; males also cooperate in aggression between communities. The chimpanzee community at Ngogo, in Kibale National Park, Uganda, is unusually large and has more males than any other known community. Field data show that adult Ngogo males groomed far more with other adult males than with females or with adolescent males, in contrast to a previous report (Ghiglieri, 1984). Adolescent males groomed adults much more than the reverse; males groomed and were groomed by females about equally. Individual males groomed mostly with a small number of other males. On average, males at Ngogo had only slightly more male grooming partners overall and had the same number of important partners as those of males in a much smaller community in the Mahale National Park, Tanzania, and they distributed their grooming less equitably. These results fit those expected if limits on available grooming time cause males to have a loyalty problem as the number of potential grooming and alliance partners increases. Despite differences in the extent and equitability of their grooming networks, males at both Ngogo and Mahale showed reciprocity in grooming. Grooming reciprocity has been demonstrated for captive chimpanzee males, but the Ngogo findings are the first demonstrations of reciprocity in wild communities.