Shifting through the forest: home range, movement patterns, and diet of the eastern chimpanzee (Pan troglodytes schweinfurthii) in Nyungwe National Park, Rwanda.

Populations of the endangered eastern chimpanzee (Pan troglodytes schweinfurthii) are declining throughout their range. Although Nyungwe National Park (NNP) harbors the largest remaining eastern chimpanzee population in Rwanda, we know little about their space use and dietary patterns. We studied home range, movement, and diet of two communities of chimpanzees in NNP using daily tracking data (6:00 am to 6:00 pm) collected from 2000 to 2015. One community, Mayebe, resided in the forest center, and the other community, Cyamudongo, inhabited a forest fragment located about 10 km from the main forest. Home range estimated with the 95% kernel density estimation (KDE) method was 21 km2 for the Mayebe community and 4 km2 for the Cyamudongo community. Chimpanzee home range sizes were smaller during the dry versus wet season and varied monthly throughout the year. The Mayebe community had an average hourly step length of 75 ± SE 5 m with a daily movement range of 987 ± SE 71 m, while the Cyamudongo community had a shorter hourly step length of 52 ± SE 3 m with a daily movement range of 651 ± SE 71 m. Both chimpanzee communities fed primarily on Ficus spp. Other important dietary items included fruits of Symphonia globulifera, Syzygium guineense, and Chrysophyllum gorungosanum for the Mayebe community and Trilepisium madagascariense for the Cyamudongo community. Food choice varied monthly and seasonally for each chimpanzee community. Our study provides the first estimates of home range size and movement parameters for chimpanzees in Rwanda and documents their food habits and seasonal variations therein. We also identified the 50% core home range for each chimpanzee community and suggest this area as the focus of management actions. These results could help park management reduce threats to chimpanzees and other sympatric species by improving the efficiency of ranger patrols.

The genetic population structure of wild western lowland gorillas (Gorilla gorilla gorilla) living in continuous rain forest.

To understand the evolutionary histories and conservation potential of wild animal species it is useful to assess whether taxa are genetically structured into different populations and identify the underlying factors responsible for any clustering. Landscape features such as rivers may influence genetic population structure, and analysis of structure by sex can further reveal effects of sex-specific dispersal. Using microsatellite genotypes obtained from noninvasively collected fecal samples we investigated the population structure of 261 western lowland gorillas (WLGs) (Gorilla gorilla gorilla) from seven locations spanning an approximately 37,000 km(2) region of mainly continuous rain forest within Central African Republic (CAR), Republic of Congo and Cameroon. We found our sample to consist of two or three significantly differentiated clusters. The boundaries of the clusters coincided with courses of major rivers. Moreover, geographic distance detoured around rivers better-explained variation in genetic distance than straight line distance. Together these results suggest that major rivers in our study area play an important role in directing WLG gene flow. The number of clusters did not change when males and females were analyzed separately, indicating a lack of greater philopatry in WLG females than males at this scale.

Lethal combats in the forest among wild western gorillas.

Lethal intergroup encounters occur in many species because of sexual selection. While documented in mountain gorillas, they are absent in western gorillas as, instead, it is predicted by their higher feeding (frugivory) and mate competition (single-vs. multi-male groups). We investigate whether the injuries on three dead silverbacks and one adult female from four groups of western gorillas in the Central African Republic, resulted from interactions with gorillas or leopards. We identified two distinct injury patterns caused by gorillas (isolated lacerations, round wounds) and leopards (punctures clustered on head/neck) by analyzing injuries caused by mountain gorillas and leopards to gorillas and non-gorilla species, respectively. The western gorilla injury pattern is similar to that of mountain gorillas suggesting that lethal encounters occur, albeit infrequently, as predicted by sexual selection in a one-male society. While sexual dimorphism and polygynous sociality favored the evolution of violent encounters, multiple males in groups may influence their frequency.

Feeling a bit peckish: Seasonal and opportunistic insectivory for wild gorillas.

OBJECTIVES: Insectivory likely contributed to survival of early humans in diverse conditions and influenced human cognitive evolution through the need to develop harvesting tools. In living primates, insectivory is a widespread behavior and frequently seasonal, although previous studies do not always agree on reasons behind this. Since western gorillas (Gorilla gorilla) diet is largely affected by seasonal variation in fruit availability, we aimed to test three non-mutually exclusive hypotheses (habitat use, frugivory and rainfall) to explain seasonality in termite feeding across age/sex classes in three habituated groups (Nindividuals = 27) in Central Africa. MATERIALS AND METHODS: We used 4 years of ranging, scan and continuous focal sampling records of gorillas (Nranging days = 883, Nscans = 12,384; Nhours = 891) in addition to 116 transects recording vegetation and termite mound distribution. RESULTS: Depending on the age/sex classes, we found support for all three hypotheses. Time spent in termite-rich vegetation positively impacted termite consumption in all age/sex classes, but subadults. Lengthier travels increased termite feeding in females but decreased it in subadults. Frugivory decreased termite consumption in adults. Daily rainfall had a positive effect on termite feeding and foraging in silverbacks and juveniles, but a negative effect in subadults. For females, rainfall had a positive effect on termite feeding, but a negative effect for termite foraging. DISCUSSION: In great apes, seasonal insectivory seems to be multifactorial and primarily opportunistic with important differences among age/sex classes. While insectivory has potentials to be traditional, it likely played a crucial role during primate evolution (including ours), allowing diet flexibility in changing environments.

Western lowland gorillas (Gorilla gorilla gorilla) change their activity patterns in response to frugivory.

The most important environmental factor explaining interspecies variation in ecology and sociality of the great apes is likely to be variation in resource availability. Relatively little is known about the activity patterns of western lowland gorillas (Gorilla gorilla gorilla), which inhabit a dramatically different environment from the well-studied mountain gorillas (G. beringei beringei). This study aims to provide a detailed quantification of western lowland gorillas' activity budgets using direct observations on one habituated group in Bai Hokou, Central African Republic. We examined how activity patterns of both sexes are shaped by seasonal frugivory. Activity was recorded with 5-min instantaneous sampling between December 2004 and December 2005. During the high-frugivory period the gorillas spent less time feeding and more time traveling than during the low-frugivory period. The silverback spent less time feeding but more time resting than both females and immatures, which likely results from a combination of social and physiological factors. When compared with mountain gorillas, western lowland gorillas spend more time feeding (67 vs. 55%) and traveling (12 vs. 6.5%), but less time resting (21 vs. 34%) and engaging in social/other activities (0.5 vs. 3.6%). This disparity in activity budgets of western lowland gorillas and mountain gorillas may be explained by the more frugivorous diet and the greater dispersion of food resources experienced by western lowland gorillas. Like other apes, western lowland gorillas change their activity patterns in response to changes in the diet.

The Influence of Seasonal Frugivory on Nutrient and Energy Intake in Wild Western Gorillas.

The daily energy requirements of animals are determined by a combination of physical and physiological factors, but food availability may challenge the capacity to meet nutritional needs. Western gorillas (Gorilla gorilla) are an interesting model for investigating this topic because they are folivore-frugivores that adjust their diet and activities to seasonal variation in fruit availability. Observations of one habituated group of western gorillas in Bai-Hokou, Central African Republic (December 2004-December 2005) were used to examine seasonal variation in diet quality and nutritional intake. We tested if during the high fruit season the food consumed by western gorillas was higher in quality (higher in energy, sugar, fat but lower in fibre and antifeedants) than during the low fruit season. Food consumed during the high fruit season was higher in digestible energy, but not any other macronutrients. Second, we investigated whether the gorillas increased their daily intake of carbohydrates, metabolizable energy (KCal/g OM), or other nutrients during the high fruit season. Intake of dry matter, fibers, fat, protein and the majority of minerals and phenols decreased with increased frugivory and there was some indication of seasonal variation in intake of energy (KCal/g OM), tannins, protein/fiber ratio, and iron. Intake of non-structural carbohydrates and sugars was not influenced by fruit availability. Gorillas are probably able to extract large quantities of energy via fermentation since they rely on proteinaceous leaves during the low fruit season. Macronutrients and micronutrients, but not digestible energy, may be limited for them during times of low fruit availability because they are hind-gut fermenters. We discuss the advantages of seasonal frugivores having large dietary breath and flexibility, significant characteristics to consider in the conservation strategies of endangered species.

Endoparasites of western lowland gorillas (Gorilla gorilla gorilla) at Bai Hokou, Central African Republic.

A coprologic study of free-ranging western lowland gorillas (Gorilla gorilla gorilla) at Bai Hokou, Dzangha-Ndoki National Park, Central African Republic (2 degrees 51'34''N, 16 degrees 28'03''E) was conducted from October 1999 to November 2000. All 75 fecal samples examined were positive for endoparasites, and each contained at least two species. Parasites present included two genera of amoebae, entodiniomorph ciliates, including Prototapirella gorillae, Troglodytella spp., and Gorillophilus thoracatus, a Balantidium-like organism, strongyle/trichostrongyle eggs (including a presumptive Mammomonogamus sp. and several other genera), Strongyloides sp., Probstmayria sp., a spirurid, a trichuroid, and several unidentified trematodes. Flagellates and cestodes were not found. Despite the presence of a variety of parasite genera, in general, levels of parasitism were low. These data provide baseline parasitologic data for this population as part of a comprehensive health-monitoring program. With the advent of ecotourism in this study area, continued monitoring is indicated for insuring the health of both gorillas and humans in the Bai Hokou study area.

Effects of group dynamics and diet on the ranging patterns of a western gorilla group (Gorilla gorilla gorilla) at Bai Hokou, Central African Republic.

This study describes how group dynamics and diet have influenced the ranging patterns of a western gorilla group at Bai Hokou, Central African Republic. The results are compared with those from an earlier study [Cipolletta, International Journal of Primatology, 2003], when the same group was larger and undergoing the process of habituation to humans. Data were obtained from maps of the gorillas' travel routes, direct observations, and analysis of fecal samples. Through the years, the group has experienced a decrease in size, from eight to three individuals, with periods of membership fluctuation. The male's search for new mates resulted in a larger home range than was recorded when the group consisted of more individuals. Moreover, despite an average group size of three throughout this study, the monthly range and mean daily path length (DPL) were also larger when the group was acquiring/losing members in new areas, than when no new members joined or left the group. Fruit was consumed year-round, although more heavily so during wet months. The influence of fruit consumption on the ranging patterns was concealed initially by the effect of habituation [Cipolletta, International Journal of Primatology, 2003], and later (at least partially) by the male's search for new mates. In the last 14 months of the study, when the group numbered only three individuals and was ranging in a restricted area, the average DPL, but not the monthly range, increased when the gorillas were consuming more fruit.

Social structure and life-history patterns in western gorillas (Gorilla gorilla gorilla).

Life-history traits and ecological conditions have an important influence on primate social systems. Most of what we know about the life-history patterns and social structure of gorillas comes from studies of eastern gorillas (Gorilla beringei sp.), which live under dramatically different ecological conditions compared to western gorillas (Gorilla gorilla sp.). In this paper we present new data on western gorilla social structure and life histories from four study sites, and make comparisons with eastern gorilla populations. Data were obtained from two study sites with gorilla groups undergoing the habituation process (Lossi, Democratic Republic of Congo and Bai Hokou, Central African Republic) and two "bai" studies (Maya Nord and Mbeli Bai, Republic of Congo). The size and structure of these groups were similar to those seen in eastern gorillas. However, differences in the occurrence of various group transitions (group formations, changes between one-male and multimale composition, and group disintegrations) exist, and western gorillas notably exhibit much higher rates of male emigration and correspondingly fewer multimale groups compared to mountain gorillas. Certain phenomena have been observed only rarely, including predation by leopards. The preliminary data show no significant differences in birth rates between western gorillas and mountain gorillas. The ecological variability across gorilla habitats likely explains the flexibility in the social system of gorillas, but we need more information on the social relationships and ecology of western gorillas to elucidate the causes for the similarities and differences between western and eastern gorillas on the levels of individuals, social groups, and population dynamics.

Western gorilla diet: a synthesis from six sites.

The objective of this paper is to collate information on western gorilla diet from six study sites throughout much of their current range, including preliminary information from two sites (Afi and Lossi), where studies of diet have begun only recently. Food lists were available from each site, derived from indirect signs of gorilla feeding (such as feces), with some observational data. Important staple, seasonal, and fallback foods have been identified, and a number of striking similarities across sites have been revealed based on a much larger data set than was previously available. It was confirmed that the western gorilla diet is always eclectic, including up to 230 items and 180 species. The greatest diversity is found among the fruit species eaten, fruit being included in western gorilla diets from all sites and throughout most or all of the year. Eight plant families provide important foods at five, or all six, sites, suggesting that it may be possible in the future to predict which habitats are the most suitable for gorillas. Gorillas exploit both rare and common forest species. Similarities and differences among sites can be explained superficially on the basis of geography and the past history of the forest. Gorilla density across sites appears to be most affected by the density of monocotyledonous bulk food plants, but its relationship to the density of important tree food species has yet to be tested.