Food Supply and Chimpanzee (Pan troglodytes schweinfurthii) Party Size in the Budongo Forest Reserve,Uganda

A central issue in socioecology is the nature of the relationship between an organism's environment and its social structure. In chimpanzees, the fission-fusion social system is thought to minimize feeding competition for primary dietary components: ephemeral, dispersed patches of ripe fruit. Intragroup feeding competition is thought to force individuals into small parties. Informal observations in the Sonso region of the Budongo forest had suggested that in this habitat, food supply was such that feeding competition was less important in determining grouping patterns than elsewhere. We used data collected on food supply and party sizes over a 4-year period to investigate this suggestion. In accord with theoretical expectation, sizes of foraging parties fluctuated with the size of food patches. However, party sizes showed either negative or no relationship with habitat-wide measures of food abundance. Likewise party sizes showed little relationship to overall measures of food dispersion. For important dietary items, both fruit and leaves had patchy distributions, though the degree of clumping was not strong, and fruit was not more clumped than leaves. Generally, abundant food appeared to be less patchy, and chimpanzees appeared to use more patches as food became more abundant rather than forming larger parties. We suggest that both dispersal and abundance need to be considered when investigating the impact of food supply on grouping patterns, and that the importance of food as a factor in determining chimpanzee grouping patterns declines with increasing levels of abundance.     

Effect of Group Size on Activity Budgets of Colobus vellerosus in Ghana

Group size influences foraging efficiency in several primates. We examined the activity budgets of 3 groups of Geoffroy's pied colobus (Colobus vellerosus) at the Boabeng-Fiema Monkey Sanctuary in Ghana to determine whether larger group size induces scramble competition. We studied 2 groups (B1; N = 7-8 and WW; N = 31-33) occupying slightly overlapping home ranges from August to November 2000. We observed the third group, B2 (N = 15-16), comprising B1 and 7 male invaders in the same home range as B1 from August to November 2001. By comparing groups belonging to the same population and occupying sligthly overlapping or similar home ranges, we were able to control, to a certain extent, for differences in food distribution. We recorded a total of 3353 scans, yielding 14,886 activity records, over 73 days. As with other black-and-white colobus, resting was their most common activity (59%). Intergroup comparisons suggest that time spent feeding, resting and moving did not vary in relation to group size. However, intragroup comparisons between the sexes show that females in the large group spent more time feeding than males did, whereas this was not the case in the small group, which suggests that scramble competition may be occurring among female Colobus vellerosus at BFMS. It is also possible that this may be due to greater nutritional requirements because of a higher proportion of infants in the large group. In fact, the proportion is quite similar between the two groups, lending support to the idea that females in the two groups had comparable nutritional demands due to lactation. This suggests that increased feeding in females in the large group was partly an effect of scramble competition. Group size and group composition also influenced the frequency of social behavior. There was more grooming in the large group, and it was performed mostly by females. The distribution of activities throughout the day was similar to the pattern reported for other black-and-white colobus.     

Dominance style in female guerezas (Colobus guereza Rüppell 1835)

Socioecological models assume that quality and distribution of food ultimately determine female social relationships: a high quality diet and clumped food distribution should result in the establishment of a hierarchy with stable rank relations which is supported by empirical studies on frugivorous cercopithecines. By contrast, folivorous species with their low quality diet and dispersed food distribution should have egalitarian social relationships but empirical data are very rare. This study on female guerezas of a zoo group aimed to test the models in a colobine species and the results largely agreed with the predictions of the models: facial expressions, vocalizations, and gestures were not used for signalling dominance or subordination. Unritualized aggressions occurred frequently but were of low intensity, and interventions by third parties were never observed. Aggressions were exchanged bidirectionally and this was true also for food stealing and retreats. All this indicated the lack of established rank relations. Allogrooming was distributed rather equally and showed no kin bias. All these features characterize egalitarian social relationships and, hence, support the socioecological models.     

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.     

Plant Species with Potential as Food,Nesting Material,or Tools at a Chimpanzee Refuge Site in Caddo Parish,Louisiana

Surplus chimpanzees live in research laboratories where they will likely remain for the rest of their lives. An alternative to laboratory housing is an outdoor enclosure in a warm climate. Before construction, researchers should conduct a vegetation survey because chimpanzees use vegetation daily. Chimp Haven, Inc. is developing a chimpanzee sanctuary in Caddo Parish, Louisiana, and we conducted a vegetation survey at the site to 1) characterize the abundance and structure of vegetation; 2) identify plants with utility as food, nesting material, or tools; 3) determine the abundance and distribution of useful vegetation; and 4) identify any possibly hazardous vegetation. We established 48 plots, quantified the woody vegetation, and identified potentially useful and hazardous plant species from the published literature. Vegetation at the site was secondary growth from logging of loblolly pine (Pinus taeda), the most abundant tree. Other abundant trees included oak (Quercus spp.), elm (Ulmus spp.), sweetgum (Liquidambar styraciflua), and maple (Acer spp.). Small plants (1–9.9 m tall) constituted 95% of all vegetation. Forty-one of 64 plant species have edible vegetative parts, while 35 species have edible reproductive parts. We identified 55 potential nesting trees by size (dbh ≥ 25 cm) and 4 species used as tools by captive great apes. Useful vegetation was concentrated in areas with topographic relief or in a floodplain, while flat areas with fields had less. We recommend that the floodplain be cropped for fruit and browse and enclosures encompass areas with relief and fields, creating a diverse habitat.     

Effects of Group Size on Ranging Patterns in Brachyteles arachnoides hypoxanthus

We collected systematic data on the home range and day ranges of one group of 57–63 muriquis (Brachyteles arachnoides hypoxanthus) at the Estação Biológica de Caratinga, Minas Gerais, Brazil from September 1998–July 1999, and compared them with similar data collected 15 years ago when the 23–27 individuals in the group traveled together as a cohesive unit. Home range size increased from 168 ha to 309 ha, reflecting an expansion into areas of the forest that were previously unutilized and consistent with the positive relationship predicted between group size and home range size. By contrast, muriquis exhibited remarkable seasonal and interannual stability in their day ranges. Day ranges, which were calculated from 144 days with 8 h of observation, averaged 1,313 ± 573 m (median = 1,206 m). Day ranges did not vary with the size of subgroups, defined as independent individuals that traveled with one another out of contact with other group members. Subgroups were significantly larger during the rainy season (mean = 41.8 ± 12.7, median = 46.0 individuals, n = 72) than the dry season (mean = 36.6 ± 13.25, median = 39.5 individuals, n = 72). Subgroups were also larger than the size of the entire group during the previous study, yet their day ranges are indistinguishable. The stability in muriqui day ranges is consistent with predictions for folivorous primates in which other indicators of intragroup feeding competition, such as female dominance relationships, are also absent. We attribute the transition from cohesive to fluid grouping patterns to limits on the number of individuals that can coordinate their movements when they spread out while foraging and suggest that seasonal differences in subgroup sizes without corresponding adjustments in day ranges reflect seasonal differences in the distribution of preferred foods coupled with the effects of reproductive seasonality on muriqui grouping patterns.     

Chimpanzee predation in the Mahale mountains from August 1979 to May 1982

Fifty-four episodes of predatory behavior of wild chimpanzees were recorded in Mahale, western Tanzania, from August 1979 to May 1982. The chimpanzees most frequently hunt in two seasons, during May, and from August to December. Longer-term fecal analysis indicates that predation frequency is significantly higher in the dry than in the rainy season. The seasonality of predation might be the result of the sum of various ecological factors, at least one of which is the birth season of the prey species. Most of the prey are juvenile blue duiker, bushbuck, bushpig, red colobus, and red-tailed monkeys. Sex difference is recognized in the prey selection and in the hunting method employed. Apparent local difference in the predatory behavior between Mahale and Combe chimpanzees (in Mahale,females hunt more frequently, and blue duiker is the most frequent prey) can be understood in terms of the difference either in the observation methods or in the faunal diversity and density. Other aspects of predatory behavior also are reported.     

Group Size, Home Range Use, and Seasonal Variation in the Ecology of Eulemur mongoz

We collected data during a 10-month study carried out on the mongoose lemur, Eulemur mongoz, at Anjamena in northwestern Madagascar, which provide baseline information on seasonal variation in the ecology, home range use and some aspects of the behavior of two neighboring groups. We monitored group size of nine groups in the study area and assessed them for seasonal variation. We present additional information collected during short-term surveys in other areas before and during the study for comparison. The study groups were small family units, and changes in group size were limited to births and emigrations of sexually mature progeny. In spite of clear seasonal changes in climate and vegetation, there is no variation in grouping patterns, so it is not possible to correlate variation in group size with seasonal variation of ecological variables. Comparison with ecological data from other field studies on lemurids reveals differences in food resource distribution in western forests versus other types of Malagasy forest. This distribution of food resources may predict home range size in mixed frugivorous–folivorous lemurs. Small home ranges, mainly in the West, could be correlated with a uniform distribution of food resources. Finally, we suggest that the dry season in the West may not present frugivorous–folivorous lemurs with major problems in finding an adequate food supply. This is supported by the lack of seasonal differences in ranging behavior of mongoose lemurs.     

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.     

Seminomadic Ranging in a Population of Black Colobus (Colobus satanas) in Gabon and its Ecological Correlates

We observed ranging patterns in a black colobus population in the Forêt des Abeilles, Central Gabon. We compared the results with those for two other populations (Lopé Reserve in Central Gabon, Douala-Edéa in coastal Cameroun) in order to estimate the extent of specific variability in population structure and ranging behavior and to identify ecological factors influencing interpopulation differences. The mean number of monkeys in a group is 17, and all groups had a multimale structure. Home ranges were large, and the fact that they continued to increase with increasing number of observations indicated that the monkeys did not frequently resample their habitat. Ranging patterns varied according to the staple food consumed: distances travelled daily increased with increasing seed intake and decreased with increasing leaf intake. The ranging patterns of groups of Colobus satanas are seminomadic. Up to 6 or 7 groups shared the same space, and two group ranges could overlap by as much as 65–75%. Comparisons with the two populations previously studied showed that group size and group structure are broadly similar. At all three sites, black colobus were mainly seed eaters and ate large amounts of leaves in the season of fruit shortage. In the Forêt des Abeilles, however, group home ranges were much larger than at other sites, and this population had the lowest density. This is correlated with vegetation composition and with harshness of the main dry season. This colobus population appears limited by both recurring food shortage in the dry season and episodic periods of seed shortage resulting from irregular fruiting of the dominant family Caesalpiniaceae. Their seminomadic ranging would constitute the least costly strategy to cope with the low carrying capacity of their habitat.     

Chimpanzee microsatellite PCR primers applied to paternity testing in a captive colony

Previously designed primers for the polymerase chain reaction (PCR) amplifying microsatellite DNA segments containing GT/AC dinucleotide repeats in the chimpanzee (Pan troglodytes) genome were used for paternity testing in a breeding colony in captivity. Combinations of three PCR primers identified the fathers of all the tested 40 chimpanzees born in an eight-year period. The results suggested: (1) a positive (though not conclusive) correlation between male rank and number of offspring; (2) choice of mating partners by the female rather than by the male; and (3) absence of stable mating pairs over the years. For studies of chimpanzees in captivity and in the wild, these primers should be useful for paternity testing, for investigating genetic variations, and for improving genetic maintenance of breeding colonies.     

Behavioral Ecology of Sympatric Chimpanzees and Gorillas in Bwindi Impenetrable National Park,Uganda: Diet

Via a field study of chimpanzees (Pan troglodytes schweinfurthii) and gorillas (Gorilla gorilla beringei) in Bwindi Impenetrable National Park, Uganda, we found that their diets are seasonally similar, but diverge during lean seasons. Bwindi chimpanzees fed heavily on fruits of Ficus sp., which were largely ignored by the gorillas. Bwindi gorilla diet was overall more folivorous than chimpanzee diet, but was markedly more frugivorous than that of gorillas in the nearby Virunga Volcanoes. During 4 mo of the year Bwindi gorilla diet included more food species than that of the chimpanzees. Three factors in particular—seasonal consumption of fibrous foods by gorillas, interspecific differences in preferred fruit species, and meat consumption by chimpanzees—contributed to dietary divergence between the two species. When feeding on fruits, gorillas ate Myrianthus holstii more frequently than chimpanzees did, while chimpanzees included more figs in their annual diet. Chimpanzee diet included meat of duikers and monkeys; gorilla frequently consumed decaying wood.     

Feeding,Diet, and Jaw Form in West African Colobus and Procolobus

The functional link between mandibular morphology and masticatory stress has been documented by both experimental and comparative investigation. Somewhat more tenuous is the purported connection between dietary variation and the form of the jaws in primates. Several factors complicate the inference of such a connection, including anecdotal or incomplete dietary data from field studies and allometric effects on skeletal form that may have little to do with diet per se. We compared the jaws of sympatric colobines from West Africa to test the effect of diet on mandibular form. Procolobus badius and Colobus polykomos occupy the same habitat yet differ in diet primarily due to the exploitation of hard seeds by C. polykomos. The fact that the two taxa are comparable in body size also obviates the need for allometric qualifications. Colobus polykomos is expected to possess more robust mandibular corpora than Procolobus badius. In fact, the jaws of Colobus polykomos do not differ consistently from those of Procolobus badius in terms of biomechanical function. This apparent failure of mandibular morphology to reflect differences in diet and feeding behavior may be due to a variety of factors. We suspect that functional demands related to canine tooth support are contributing to obliteration of the expected biomechanical signal. Successful prediction of dietary effects on mandibular form requires consideration of competing structural and functional demands. The influence of diet on mandibular corporeal morphology is not equivalent across different primate species.     

Jane goodall and the chimpanzees of gombe: An analysis of publications and their impact on teaching science

Five types of introductory university textbooks (N=37) were analyzed for references and citations pertaining to research on wild chimpanzees. Jane Goodall's publications were cited about three times as often as the publications from field sites other than Gombe and approximately five times more often than other Gombe researchers. Biological anthropology textbooks cited Goodall's work most often, followed by textbooks in general anthropology and cultural anthropology. Psychology and biology textbooks cited Goodall least often. Goodall's most comprehensive work, The Chimpanzees of Gombe: Patterns of Behavior (1986), was the most often cited publication about Gombe's apes, and tool-use was the most cited topic. The number of citations to wild chimpanzees tripled from publications in the 1960s to those in the 1980s, suggesting a growing recognition of primatology in the teaching of science.     

Chimpanzee tool technology in the Goualougo Triangle, Republic of Congo

With the exception of humans, chimpanzees show the most diverse and complex tool-using repertoires of all extant species. Specific tool repertoires differ between wild chimpanzee populations, but no apparent genetic or environmental factors have emerged as definitive forces shaping variation between populations. However, identification of such patterns has likely been hindered by a lack of information from chimpanzee taxa residing in central Africa. We report our observations of the technological system of chimpanzees in the Goualougo Triangle, located in the Republic of Congo, which is the first study to compile a complete tool repertoire from the Lower Guinean subspecies of chimpanzee (Pan troglodytes troglodytes). Between 1999 and 2006, we documented the tool use of chimpanzees by direct observations, remote video monitoring, and collections of tool assemblages. We observed 22 different types of tool behavior, almost half of which were habitual (shown repeatedly by several individuals) or customary (shown by most members of at least one age-sex class). Several behaviors considered universals among chimpanzees were confirmed in this population, but we also report the first observations of known individuals using tools to perforate termite nests, puncture termite nests, pound for honey, and use leafy twigs for rain cover. Tool behavior in this chimpanzee population ranged from simple tasks to hierarchical sequences. We report three different tool sets and a high degree of tool-material selectivity for particular tasks, which are otherwise rare in wild chimpanzees. Chimpanzees in the Goualougo Triangle are shown to have one of the largest and most complex tool repertoires reported in wild chimpanzee populations. We highlight new insights from this chimpanzee population to our understanding of ape technological systems and evolutionary models of tool-using behavior.     

Comparison of habitat quality and diet of Colobus vellerosus in forest fragments in Ghana

The forest fragments surrounding the Boabeng-Fiema Monkey Sanctuary (BFMS) in central Ghana shelter small populations of Colobus vellerosus. Little is known about these populations or the ability of the fragments to support them, despite the fact that these fragments represent potentially important habitat for the colobus in this region. We compared the diet of three groups of C. vellerosus in the fragments to two groups in BFMS. We also examined the differences in plant species composition and food abundance among fragments. The study took place from June to November 2003. Dietary data were collected using scan sampling. Plant species composition and food abundance were evaluated using tree plots and large tree surveys. As in BFMS groups, leaves constituted the highest proportion of the diet of fragment groups, yet the colobus in fragments fed on more lianas than did those in BFMS. Over 50% of all species observed eaten by colobus in the fragments were not consumed in BFMS groups during the same season. Food abundance was similar between fragments and BFMS, although species composition differed. There was no relationship between the density of colobus and the density of food trees or percentage of food species, suggesting that other factors may be influencing the number of colobus present. This study highlights the broad dietary range of C. vellerosus, which may be a factor allowing its survival in these fragments.     

Noninvasive Monitoring of the Health of <Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis> in the Kibale National Park,Uganda

We assessed the health status of chimpanzees (Pan troglodytes schweinfurthii) of the Kanyawara group in the Kibale National Park in Western Uganda via noninvasive methods. We conducted visual veterinary inspection, parasitological and urine analysis in association with behavioral observations, causing minimal disturbance or stress to individually recognized chimpanzees. We applied multiple parasitological techniques to 252 stool samples to compare their efficacy in detecting parasitic infection and to increase the power of detecting a wide range of parasites at a more sensitive level. We examined 76 urine samples via a quick detection method to evaluate multiple parameters of urine that indicate organ dysfunction. Results of the different analyses are mutually supportive and provided useful information for monitoring bodily condition and diseases. The multifaceted health evaluation system is a beneficial tool for monitoring long-term and short-term changes in health status due to environmental stress, seasonal dietary change, and disease in wild chimpanzee populations. Use of this method to detect changes in health, when employed together with behavioral observations, may also provide important insights into the potential effects of self-medicative behaviors.