Western lowland gorilla diet and resource availability: new evidence,cross-site comparisons,and reflections on indirect sampling methods

We describe the resource availability and diet of western lowland gorillas (Gorilla gorilla gorilla) from a new study site in the Central African Republic and Republic of Congo based on 3 years of study. The results, based on 715 fecal samples and 617 days of feeding trails, were similar to those reported from three other sites, in spite of differences in herb and fruit availability. Staple foods (consumed year-round) included high-quality herbs (Haumania), swamp herbs (when present), and a minimal diversity of fruit. A variety of fruits (average of 3.5 species per day and 10 per month) were selectively consumed; gorillas ignored some common fruits and incorporated rare fruits to a degree higher than predicted based on availability. During periods of fruit abundance, fruit constituted most of the diet. When succulent fruits were unavailable, gorillas used low-quality herbs (i.e., low-protein), bark, and more fibrous fruits as fallback foods. Fibrous fruit species, such as Duboscia macrocarpa and Klainedoxa gabonensis, were particularly important to gorillas at Mondika and other sites as fallbacks. The densities of these two species are similar across sites for which data are available, in spite of major differences in forest structure, suggesting they may be key species in determining gorilla density. No sex difference in diet was detected. Such little variation in western lowland gorilla diet across sites and between sexes was unexpected and may partly reflect limitations of indirect sampling.     

Western lowland gorilla density and nesting behavior in a Gabonese forest logged for 25 years: implications for gorilla conservation

The tropical forests of the Congo Basin constitute biodiversity refuges that still hold large numbers of species, including endemic and endangered vertebrates. Along with several key species, the critically endangered western lowland gorilla (WLG) potentially contributes to forest dynamics through seed dispersal. Considering the extensive influence of timber harvesting on tropical forest ecosystems, the survival of gorilla populations in logged forests might prove critical for forest ecosystem conservation. We estimated WLG density, through a nest count survey, in a forest in southeast Gabon that has been logged for 25 years. Nesting behavior and habitat use were described and we applied generalized linear models to identify the factors that influence gorilla day and night habitat use. The estimated density of weaned gorillas, 1.5 gorillas km^?2, is comparable with estimates from some protected areas and other sustainably managed sites within their range. Habitat type had the greatest influence on nest site distribution. We observed a preference for nesting in open terra firma forest, and open habitats in general, which supports the findings of previous studies. Habitat use during the day was strongly influenced by habitat type and human activities, and to a lesser degree by functional and non-functional roads, and rivers. Our results support the suggestion that logged forests are suitable habitats for WLG if hunting and poaching are controlled. We recommend collaborations between timber operators and scientists to improve the conservation potential of tropical forests and enhance the wildlife-management aspects of logging practices.     

Census of gorillas in northern Republic of Congo

We censused gorilla populations in northern Congo from February to April 1989 and June 1990. The objective was to provide the first quantitative data from a variety of sites on gorilla populations from a country that had unknown but potentially high populations. The method used was a census of nests along strip transects. A total of 401.0 km of transects was sampled in four different study areas. The overall density calculated for all transects was 0.4 nesting gorillas/km². The highest density, 1.2 nesting gorillas/km², was found in the vast Likouala swamp area of north central Congo. The two northern sites showed low densities of 0.1 and 0.2 nesting gorillas/km², respectively. The northwestern site showed an intermediate density of 0.6 nesting gorillas/km². The vegetation type with the highest density was swamp forest with 2.4 nesting gorillas/km². The limited sample presented shows that gorillas are widespread and common in northern Congo, even in the swamp forests previously considered unsuitable as gorilla habitat. It is probable that Congo holds the largest population of gorillas in Africa after Gabon. © 1992 Wiley-Liss, Inc.     

Densities,Distributions, and Seasonal Movements of Gorillas and Chimpanzees in Swamp Forest in Northern Congo

The conservation of gorillas (Gorilla spp.) and chimpanzees (Pan troglodytes) depends upon knowledge of their densities and distribution throughout their ranges. However, information about ape populations in swamp forests is scarce. Here we build on current knowledge of ape populations by conducting line transect surveys of nests throughout a reserve dominated by swamp forest: the Lac Télé Community Reserve in northern Congo. We estimated gorilla and chimpanzee densities, distributions across habitats, and seasonal changes in abundance. Gorilla density was 2.9 gorillas km^–2, but densities varied by habitat (0.3–5.4 gorillas km^–2) with highest densities in swamp forest and terra firma mixed forest. Average chimpanzee density is 0.7 chimpanzees km^–2 (0.1–1.3 chimpanzees km^–2), with highest densities in swamp forest. Habitat was the best predictor of ape nest abundance, as neither the number of human indices nor the distance from the nearest village predicted nest abundance. We recorded significantly greater numbers of apes in terra firma forest during the high-water season than the low-water season, indicating that many gorillas and chimpanzees are at times concentrated in terra firma forest amid a matrix of swamp forest. Seasonally high numbers of apes on terra firma forest islands easily accessible to local people may expose them to substantial hunting pressure. Conversely, the nearly impenetrable nature of swamp forests and their low value for logging makes them promising sites for the conservation of apes.     

Plant selection for nest building by western lowland gorillas in Cameroon

We examined 834 nests built by western lowland gorillas in Cameroon between July 2008 and July 2011 to identify the plant species used in their construction. Preference for each plant species for nesting was assessed using a ‘preference index’ calculated by combining information on the occurrence of each species in the forest and in the nests. Forty-six plant species representing about 15 % of the total number of species in the forest and 26 % of species used for nest building were frequently used by gorillas. Preference levels significantly varied among these species. Nests were mostly built with herbs of the families Marantaceae and Zingiberaceae and woody species such as Manniophyton fulvum (liana) and Alchornea floribunda (shrub). As observed in other gorilla populations, suitability for nest building and availability of gorilla food in stems were the likely determinants of plant selection. The total number of species used per nest ranged from 1 to 11, with an average of 4.9. This is high compared to other sites, emphasizing variability in the availability of nest building materials and habitat differences across the range of the western gorilla. Seasonal changes in the use of different habitat types for nesting did not appear to influence plant use for nest building as there was little variation in plant selection across seasons or the composition of nests. Our findings suggest that gorillas non-randomly select plant species to build nests, and use a particular set of species combined at varying proportions, with no clear seasonal or spatial patterns.     

Sympatric Chimpanzees (<Emphasis Type="Italic">Pan troglodytes troglodytes</Emphasis>) and Gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>) in Loango National Park,Gabon: Dietary Composition,Seasonality, and Intersite Comparisons

Dietary overlap of sympatric apes is complex and understudied, but its examination is essential to further our understanding of species distribution, abundance, and community ecology. Over 3 yr we studied food availability and dietary composition of central chimpanzees (Pan troglodytes troglodytes) and western gorillas (Gorilla gorilla gorilla) in Loango National Park, Gabon. We predicted that living in a habitat dominated by mature forest with sparse terrestrial herbaceous vegetation would lead to an increase in frugivory by gorillas, resulting in increased dietary overlap between the 2 ape species vs. other sites, but that chimpanzees would remain more frugivorous than gorillas. Through fecal analysis we measured overlap in fruit consumption between the 2 species on a bimonthly basis using the Renkonens method. Mean overlap was 27.5% but varied greatly seasonally, ranging between 0.3% and 69%, indicating that when examined on a finer scale, the degree of overlap appears much lower than at other study sites. In contrast to studies elsewhere, there was not a positive correlation between rainfall and fruit availability in Loango, and the long dry season was a period of high fruit production. As observed elsewhere, we found a positive correlation between fruit consumption and fruit availability for both chimpanzees and gorillas and we found that chimpanzees were more frugivorous than gorillas. A very low availability of herbs did not lead to increased frugivory by gorillas nor increased overlap between the 2 ape species vs. other field sites. We conclude that forest composition, fruit availability, and dietary variability of sympatric species can vary greatly between locations, and that chimpanzees and gorillas can adapt to primary forest with little undergrowth, where they concentrate their diet on fruit and leaves.     

Impact of ecological and social factors on ranging in western gorillas

We examined the influence of ecological (diet, swamp use, and rainfall) and social (intergroup interaction rate) factors on ranging behavior in one group of western gorillas (Gorilla gorilla gorilla) during a 16-month study. Relative to mountain gorillas, western gorillas live in habitats with reduced herb densities, more readily available fruit (from seasonal and rare fruit trees), and, at some sites, localized large open clearings (swamps and "bais"). Ranging behavior reflects these ecological differences. The daily path length (DPL) of western gorillas was longer (mean=2,014 m) than that of mountain gorillas, and was largely related to fruit acquisition. Swamp use occurred frequently (27% of days) and incurred a 50% increase in DPL, and 77% of the variation in monthly frequency of swamp use was explained by ripe fruit availability within the swamp, and not by the absence of resources outside the swamp. The annual home-range size was 15.4 km2. The western gorilla group foraged in larger areas each month, and reused them more frequently and consistently through time compared to mountain gorillas. In contrast to mountain gorillas, intergroup encounters occurred at least four times more frequently, were usually calm rather than aggressive, and had no consistent effect on DPL or monthly range size for one group of western gorillas. High genetic relatedness among at least some neighboring males [Bradley et al., Current Biology, in press] may help to explain these results, and raises intriguing questions about western gorilla social relationships.     

Feeding Ecology of Cross River Gorillas (Gorilla gorilla diehli) at Mawambi Hills: The Influence of Resource Seasonality

Determining the composition of primate diet and identifying factors that affect food choice are important in understanding habitat requirements of primates and designing conservation plans. We studied the diet of Cross River gorillas (Gorilla gorilla diehli) in relation to availability of food resources, in a semideciduous lowland forest site (Mawambi Hills) in Cameroon, from November 2009 to September 2011. Based on 109 d of feeding trail data, 203 fecal samples, and 22 mo of phenological monitoring, we determined that gorillas consumed a total of 242 food items, including 240 plant items from 186 species and 55 taxonomic families. Mawambi gorillas diversified fruit consumption when fruit availability increased, and consumed more fibrous foods (pith, leaf, bark) during times of fruit scarcity, consistent with results of other gorilla studies. However, fruit availability was not related to rainfall, and the period of fruit scarcity was more pronounced at Mawambi than at other gorilla study sites, due to a single long dry season and extreme rainfall at the end of the rainy season that delayed fruit production and ripening. We found no relationship between the daily path length of the gorillas and fruit consumption. We found feeding habits of Mawambi gorillas to be notably similar to those of a population of Cross River gorillas at Afi Mountain, Nigeria, although subtle differences existed, possibly due to site-specific differences in forest composition and altitude. At both sites the liana Landolphia spp. was the single most important food species: the leaves are a staple and the fruits are consumed during periods of fruit scarcity. Snails and maggots were consumed but we observed no further faunivory. We suggest that tree leaves and lianas are important fallback food sources in the gorilla diet in seasonally dry forests.     

Feeding ecology of western lowland gorillas in the Nouabalé-Ndoki National Park,Congo

The feeding ecology of western lowland gorillas (Gorilla gorilla gorilla) living in the Nouabalé-Ndoki National Park, northern Congo, was surveyed for one full year. This is the first record to make clear the seasonal changes in the feeding habits of gorillas in a whole year, living in the primary lowland forest almost completely undisturbed. Fecal contents, feeding traces, and direct observation were analyzed with reference to a fruit availability survey. Although the gorillas fed largely on fruits in the forest, their basic diet was fibrous parts of plants, including shoots, young leaves, and bark. Terrestrial herbaceous vegetation, such as monocotyledons of the Marantaceae and aquatic herbs having much protein content and minerals, were frequently eaten even in the fruiting season. As these highly nutritious fibrous foods were superabundant all year, the major foods of the Ndoki gorillas seemed to be those plants. However, they selected fruits as their alternative food resources in the fruiting season. Gorillas foraged on many fruit species, while showing strong preferences for some particular species. The swamp forest, including marshy grasslands, was an important and regular habitat for the Ndoki gorillas.     

Nationwide census of gorilla (gorilla g. gorilla) and chimpanzee (Pan t. troglodytes) populations in Gabon

A census was made of gorilla and chimpanzee populations throughout Gabon between December 1980 and February 1983. The aim of the census was to estimate the total numbers of both species and describe their distributions. The method was based on nest counts from line transects which allowed the calculation of population densities of all individuals except suckling infants. Fifteen types of habitat were recognized and defined in terms of their structural features. In the initial phase of the study we did transects in each habitat-type and computed mean densities for each species in each habitat-type. In the second phase of the study we estimated the sizes of gorilla and chimpanzee populations throughout the country by extrapolation from these population density values. We did transects in all areas of the country and conducted interviews to check the accuracy of the population totals obtained by extrapolation. Corrections were made to the extrapolated totals to take into account different levels of hunting pressure and other human activities found to modify ape population densities. Total populations of 34,764 gorillas and 64,173 chimpanzees were estimated. An error of ± 20% was associated with the estimated population totals, which allows the conclusion that Gabon contains 35,000 ± 7,000 gorillas and 64,000 ± 13,000 chimpanzees. The figure for gorillas is much larger than previous estimates. This seems to be because (1) gorillas occur in almost all types of forest and are not restricted to man-made secondary forest as had been though; and (2) the geographical distribution of gorillas in Gabon is wider than previously believed. Gabon's large areas of undisturbed primary forest offer exceptional potential for conservation, not only of gorillas and chimpanzees, but also of the intact tropical rain forest ecosystems which they inhabit.     

Ape Abundance and Habitat Use in the Goualougo Triangle, Republic of Congo

Chimpanzee (Pan troglodytes troglodytes) and western gorilla (Gorilla gorilla gorilla) populations in central Africa are rapidly declining as a result of disease epidemics, commercial bushmeat hunting, and habitat destruction. Our main objective was to estimate the absolute abundance and habitat utilization of chimpanzees and gorillas in the intact forests of the Goualougo Triangle in the Republic of Congo, and in an adjacent area in which selective logging will take place in the near future. The estimates provide a unique baseline for apes inhabiting an undisturbed environment. A second objective was to compare estimates of abundance and patterns of habitat utilization generated by different techniques: 1) distance sampling of individual ape nests and nest sites along line transects, 2) direct observations of apes during reconnaissance surveys, and 3) observations of ape traces during reconnaissance surveys. We completed a total of 222 km of line transect surveys in 4 sampling areas, resulting in overall density estimates of 1.53 chimpanzees/km² and 2.34 gorillas/km² from nest sites. We generated a density estimate of 2.23 chimpanzees/km² from direct observations during reconnaissance surveys of a semihabituated community in 1 of the 4 sampling areas. Habitat use profiles that nest surveys depicted on transects differed from those of direct observations and traces we encountered on reconnaissance surveys. We found the highest overall abundance of chimpanzee nests in monodominant Gilbertiodendron forest, whereas our direct observations showed that chimpanzees preferred mixed species forest. Transects that traversed the core area of the community range had the highest encounter rates of chimpanzee nests and nest sites. Gorilla nests on transects showed a preference only for mixed species forest with an open canopy, but direct observations and traces on reconnaissance surveys clearly indicated that gorillas use several habitat types. We conclude by evaluating the precision of these nest surveys and our ability to detect future trends in ape densities in the Goualougo Triangle.

Variation in behavioral traits of two frugivorous mammals may lead to differential responses to human disturbance

Human activities can lead to a shift in wildlife species’ spatial distribution. Understanding the specific effects of human activities on ranging behavior can improve conservation management of wildlife populations in human‐dominated landscapes. This study evaluated the effects of forest use by humans on the spatial distribution of mammal species with different behavioral adaptations, using sympatric western lowland gorilla and central chimpanzee as focal species. We collected data on great ape nest locations, ecological and physical variables (habitat distribution, permanent rivers, and topographic data), and anthropogenic variables (distance to trails, villages, and a permanent research site). Here, we show that anthropogenic variables are important predictors of the distribution of wild animals. In the resource model, the distribution of gorilla nests was predicted by nesting habitat distribution, while chimpanzee nests were predicted first by elevation followed by nesting habitat distribution. In the anthropogenic model, the major predictors of gorilla nesting changed to human features, while the major predictors of chimpanzee nesting remained elevation and the availability of their preferred nesting habitats. Animal behavioral traits (body size, terrestrial/arboreal, level of specialization/generalization, and competitive inferiority/superiority) may influence the response of mammals to human activities. Our results suggest that chimpanzees may survive in human‐encroached areas whenever the availability of their nesting habitat and preferred fruits can support their population, while a certain level of human activities may threaten gorillas. Consequently, the survival of gorillas in human‐dominated landscapes is more at risk than that of chimpanzees. Replicating our research in other sites should permit a systematic evaluation of the influence of human activity on the distribution of mammal populations. As wild animals are increasingly exposed to human disturbance, understanding the resulting consequences of shifting species distributions due to human disturbance on animal population abundance and their long‐term survival will be of growing conservation importance.     

Comparing ape densities and habitats in northern Congo: surveys of sympatric gorillas and chimpanzees in the Odzala and Ndoki regions

The conservation status of western lowland gorillas and central chimpanzees in western equatorial Africa remains largely speculative because many remote areas have never been surveyed and the impact of emergent diseases in the region has not been well documented. In this study, we compared ape densities and habitats in the Lokoué study area in Odzala National Park and the Goualougo Triangle in Nouabalé-Ndoki National Park in northern Republic of Congo. Both of these sites have long been considered strongholds for the conservation of chimpanzees and gorillas, but supposedly differ in vegetative composition and relative ape abundance. We compared habitats between these sites using conventional ground surveys and classified Landsat-7 ETM+ satellite images. We present density estimates via both standing-crop and marked-nest methods for the first time for sympatric apes of the Congo Basin. The marked-nest method was effective in depicting chimpanzee densities, but underestimated gorilla densities at both sites. Marked-nest surveys also revealed a dramatic decline in the ape population of Lokoué which coincided with a local Ebola epidemic. Normal baseline fluctuations in ape nest encounter rates during the repeated passages of marked-nest surveys were clearly distinguishable from a 80% decline in ape nest encounter rates at Lokoué. Our results showed that ape densities, habitat composition, and population dynamics differed between these populations in northern Congo. We emphasize the importance of intensifying monitoring efforts and further refinement of ape survey methods, as our results indicated that even the largest remaining ape populations in intact and protected forests are susceptible to sudden and dramatic declines.     

Selective logging, habitat quality and home range use by sympatric gorillas and chimpanzees: a case study from an active logging concession in Southeast Cameroon

We examined range use by great apes during logging activities and investigated associations between local variations in ape abundance and changes in the structure of the habitat or in the availability of fruits after disturbances. We carried out two annual censuses of western lowland gorilla (G. g. gorilla) and chimpanzee populations (Pan t. troglodytes) in an active logging concession in Southeast Cameroon. The results suggest that gorillas may adapt their range use to avoid most recently logged compartments, while chimpanzees appear to be more spatially resilient to logging. In our study site, selective logging affected 10% of the forest. After logging, gorillas nested in all types of vegetation, while chimpanzees nested exclusively in mixed mature forest. Fruit availability was not affected by logging and did not explain the distribution of ape nests in the study area.     

Population Density of Chimpanzees and Gorillas in the Petit Loango Reserve,Gabon: Employing a New Method to Distinguish Between Nests of the Two Species

We studied the vegetation and population density of chimpanzees (Pan troglodytes troglodytes) and gorillas (Gorilla gorilla gorilla) in the Petit Loango Reserve, Gabon. The study site consists mainly of primary and secondary forest, and it also includes patches of savanna, inundated forest, and coastal scrub. The density of herbaceous plants of Marantaceae and Zingiberaceae, which serve as important foods for great apes at other study sites, is extremely low. It was very difficult to distinguish between the nests of chimpanzees and gorillas because most gorilla nests were made in trees. We developed a new method to identify the builder of nests via a combination of field observation of nests and fecal samples, and examination of hair samples under a scanning electron microscope. Nests of chimpanzees were mainly in primary forest, and the estimated density (0.78 individual per km ² ) is higher than the average density in Gabon and comparable to the nest density in the Lopé Reserve, Gabon. Nests of gorillas were mainly in secondary forest, and their estimated density (0.21) is similar to the average next density in Gabon but lower than the gorilla nest density in the Lopé Reserve. The low density of herbaceous foods seemed to influence the density of gorillas but not the density of chimpanzees.     

Survey of gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>) and chimpanzees (<Emphasis Type="Italic">Pan troglodytes troglodytes</Emphasis>) in Southwestern Cameroon

A study on the distribution and population density of the western lowland gorilla (Gorilla g. gorilla) and the central chimpanzee (Pan t. troglodytes) was undertaken between December 1997 and August 2000 in the Campo and Maan Forests in southwestern Cameroon. The aim of this survey was to estimate the densities of the apes in different parts of the area, to assess the importance of the region for the conservation of these endangered species and to determine the influence of human activities such as logging and hunting. The survey was based on night nest counts on a total of 665.5 km of line transects. The overall density in the Campo Forest was estimated at 0.2 gorillas/km² and at 0.63–0.78 chimpanzees/km². The overall density of chimpanzees in the Maan Forest was estimated at 0.8–1 individuals/km². Gorilla density in this area was too low to allow an estimation. The highest gorilla nest density was found in secondary forest. The gorilla density in unlogged forest was significantly lower. Chimpanzees showed a clear preference for less disturbed areas. In unlogged forest, old secondary forests (logging more than 23 years ago) and areas of recent logging with large remaining patches of primary forest, significantly higher densities were calculated than inside the more heavily exploited logging concession. In areas with both logging and high hunting pressure both species were rare or even absent. The Campo Maan area is considered a very important area for the conservation of gorillas and chimpanzees. Conservation measures are urgently required to reduce the impact of logging and hunting. The creation of the Campo Maan National Park in January 2000 is an important measure to preserve the unique biodiversity in this so far hardly protected area.     

A comparison of two mountain gorilla habitats in Bwindi Impenetrable National Park, Uganda

Bwindi Impenetrable National Park, Uganda, is a small protected area (331 km²) within which there is large climatic and altitudinal variation. Therefore we compared habitat types and forest composition between two locations to investigate differences that may influence ecological conditions for large mammals, including endangered mountain gorillas. We demonstrate that there is considerable intersite variation in habitat types and spatial and temporal availability of vegetation resources of which the most apparent are the differences in species composition of plants and fruit availability. Buhoma (the lower altitude site) has a greater diversity of plant species and higher tree and shrub densities, yet has a much lower density of herbaceous vegetation eaten by gorillas compared with Ruhija. Fruit availability is significantly higher throughout the year in Buhoma compared with Ruhija. Both sites exhibit seasonal variation in fruit availability although they do not follow the same seasonal pattern, perhaps because of inherent asynchronous phenological differences amongst individual tree species. The results of this study are important because distribution and abundance of both terrestrial herbaceous vegetation and fruit resources are believed to influence aspects of feeding ecology for large mammals.     

Physical maturation,life‐history classes and age estimates of free‐ranging western gorillas—insights from Mbeli Bai,Republic of Congo

Physical maturation and life‐history parameters are seen as evolutionary adaptations to different ecological and social conditions. Comparison of life‐history patterns of closely related species living in diverse environments helps to evaluate the validity of these assumptions but empirical data are lacking. The two gorilla species exhibit substantial differences in their environment, which allows investigation into the role of increased frugivory in shaping western gorilla life histories. We present behavioral and morphological data on western gorilla physical maturation and life‐history parameters from a 12.5‐year study at Mbeli Bai, a forest clearing in the Nouabalé‐Ndoki National Park in northern Congo. We assign photographs of known individuals to different life‐history classes and propose new age boundaries for life‐history classes in western gorillas, which can be used and tested at other western gorilla research sites. Our results show that western gorillas are weaned at a later age compared with mountain gorillas and indicate slower physical maturation of immatures. These findings support the risk‐aversion hypothesis for more frugivorous species. However, our methods need to be applied and tested with other gorilla populations. The slow life histories of western gorillas could have major consequences for social structure, mortality patterns and population growth rates that will affect recovery from population crashes of this critically endangered species. We emphasize that long‐term studies can provide crucial demographic and life‐history data that improve our understanding of life‐history evolution and adaptation and help to refine conservation strategies. Am. J. Primatol. 71:106–119, 2009. © 2008 Wiley‐Liss, Inc.     

Hibernation of predatory arthropods in semi-natural habitats

Non-crop habitats provide important resources for natural enemies. Many natural enemies hibernate in non-crop habitats, from which they may colonise arable fields in the spring. Spring colonisation ensures annual repopulation of the crop with natural enemies, allowing them to keep pace with the development of pest populations. The availability of non-crop habitats can, therefore, be crucial to successful conservation biological control. We quantified the density of overwintering natural enemies near organic Brussels sprout crops in five different non-crop habitats (short grassy field margin, herbaceous field margin, herbaceous field margin under a tree line, ditch and forest). Soil and litter samples of non-crop habitats were taken at two sites. One site was located in an open agricultural landscape, the other in a landscape dominated by mixed forest. Insects belonging to Staphylinidae, Araneae, Carabidae, Coccinellidae and Dermaptera were the most abundant. Mean densities of predatory arthropods were higher in the open agricultural landscape (290 predators m⁻²) than in the forested landscape (137 predators m⁻²). Herbaceous habitat types supported the highest densities of overwintering predators (up to 400 predators m⁻²), whereas densities in the forest were lowest (10 predators m⁻²). These results indicate that herbaceous non-crop habitats are important refugia for predators and may play a vital role in conservation biological control. Handling Editor: Arne Janssen.     

Ecological Constraints on the Foraging Effort of Western Gorillas (Gorilla gorilla gorilla) at Bai Hoköu, Central African Republic

Recent studies demonstrate that western lowland gorillas incorporate much more fruit into their diet than Virunga mountain gorillas do. Very little is known, however, about how the frugivorous behavior of western gorillas influences their daily ranging behavior, which may ultimately affect social factors such as group size and structure. I examined the influence of diet and the spatiotemporal availability of plant foods on the foraging effort of nonhabituated western lowland gorilla groups during 17 months at Bai Hoköu in the Dzanga-Ndoki National Park, Central African Republic. I determined diet from indirect methods and gorilla plant food availability and spatial distribution from phenology and line transects. Daily path length gives an estimate of foraging effort and was the distance paced, following fresh gorilla trails, from morning to evening nest sites. The availability and distribution of fruit and its consumption by gorillas varied seasonally. When concentrating on fruits, gorillas traveled significantly farther (mean = 3.1 km/day) than when their diet consisted mostly of nonfruit vegetation, such as leaves and woody pith, stems, and bark (mean = 2.1 km/day). The amount of herbaceous vegetation in the diet did not vary seasonally and did not influence daily path length. The best environmental predictor of foraging effort was fruit density, or a measure combining both density and spatial pattern: coefficient of dispersion. In addition, when fruit patches were small, path length tended to increase but not significantly. Compared with results of other studies, gorillas at Bai Hoköu travel farther (mean = 2.6 km/day) than gorillas in Gabon (mean = 1.7 km/day) and five times farther than mountain gorillas in the Virungas (mean = 0.5 km/day). Increased foraging effort of gorillas in this region, especially during the fruiting season, may have profound effects on group size and structure.