Food Preferences Among Captive Western Gorillas (Gorilla gorilla gorilla) and Chimpanzees (Pan troglodytes)

I used a zoological park setting to address food preferences among gorillas (Gorilla gorilla gorill) and chimpanzees (Pan troglodytes). Gorillas and chimpanzees are different sizes, and consequently, have been traditionally viewed as ecologically distinct. Sympatric western gorillas and chimpanzees have proved difficult to study in the wild. Limited field data have provided conflicting information about whether gorillas are fundamentally different from chimpanzees in diet and behavior. Fruit eating shapes the behavior of most apes, but it is unclear whether the large-bodied gorillas are an exception to this rule, specifically whether they are less selective and more opportunistic fruit eaters than chimpanzees are. My research provides experimental observational data to complement field data and to better characterize the diets and food preferences of the African apes. During laboratory research at the San Francisco Zoological Gardens, I examined individual and specific differences in food preferences of captive gorillas and chimpanzees via experimental paired-choice food trials with foods that varied in nutritional content. During the study, I offered 2500 paired-food choices to 6 individual gorillas and 2000 additional pairs to them as a group. I also proffered 600 food pairs to 4 individual chimpanzees. Despite expectations of the implications of body size differences for diet, gorillas and chimpanzees exhibited similar food preferences. Both species preferred foods high in non-starch sugars and sugar-to-fiber ratios, and low in total dietary fiber. Neither species avoided foods containing tannins. These data support other suggestions of African apes sharing a frugivorous adaptation.     

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.     

Fallback foods and dietary partitioning among Pan and gorilla

Recent findings on the strong preference of gorillas for fruits and the large dietary overlap between sympatric gorillas and chimpanzees has led to a debate over the folivorous/frugivorous dichotomy and resource partitioning. To add insight to these arguments, we analyze the diets of sympatric gorillas and chimpanzees inhabiting the montane forest of Kahuzi-Biega National Park (DRC) using a new definition of fallback foods (Marshall and Wrangham: Int J Primatol 28 [2007] 1219–1235). We determined the preferred fruits of Kahuzi chimpanzees and gorillas from direct feeding observations and fecal analyses conducted over an 8-year period. Although there was extensive overlap in the preferred fruits of these two species, gorillas tended to consume fewer fruits with prolonged availability while chimpanzees consumed fruits with large seasonal fluctuations. Fig fruit was defined as a preferred food of chimpanzees, although it may also play a role as the staple fallback food. Animal foods, such as honey bees and ants, appear to constitute filler fallback foods of chimpanzees. Tool use allows chimpanzees to obtain such high-quality fallback foods during periods of fruit scarcity. Among filler fallback foods, terrestrial herbs may enable chimpanzees to live in small home ranges in the montane forest, whereas the availability of animal foods may permit them to expand their home range in arid areas. Staple fallback foods including barks enable gorillas to form cohesive groups with similar home range across habitats irrespective of fruit abundance. These differences in fallback strategies seem to have shaped different social features between sympatric gorillas and chimpanzees. Am J Phys Anthropol 140:739–750, 2009. © 2009 Wiley-Liss, Inc.     

Potential for Ebola transmission between gorilla and chimpanzee social groups

Over the past decade Ebola hemorrhagic fever has emerged repeatedly in Gabon and Congo, causing numerous human outbreaks and massive die-offs of gorillas and chimpanzees. Why Ebola has emerged so explosively remains poorly understood. Previous studies have tended to focus on exogenous factors such as habitat disturbance and climate change as drivers of Ebola emergence while downplaying the contribution of transmission between gorilla or chimpanzee social groups. Here we report recent observations on behaviors that pose a risk of transmission among gorilla groups and between gorillas and chimpanzees. These observations support a reassessment of ape-to-ape transmission as an amplifier of Ebola outbreaks.     

Foods consumed by sympatric populations ofGorilla g. gorilla andPan t. troglodytes in Gabon: Some preliminary data

Data on foods consumed by gorillas and chimpanzees living in primary forest in Gabon were collected, mainly by examination of the contents of feces. Gorillas ate fruit very regularly (some fruit remains were present in 97.6% of 246 fecal samples examined), in addition to leaves, stems, pith, and bark. Some fruit remains were present in all chimpanzee fecal samples examined. Mean numbers of fruit species per fecal sample were 2.5 for gorillas and 2.1 for chimpanzees. Sixty percent of all identified foods recorded for gorillas were recorded for chimpanzees as well. Our results indicate that important differences in diet exist between western lowland gorillas and the eastern gorilla populations of Kahuzi-Biega and the Virunga Volcanoes. It is now clear that western gorillas cannot be accurately classed as folivores.     

Insect-eating by sympatric Lowland gorillas (Gorilla g. gorilla) and chimpanzees (Pan t. troglodytes) in the Lopé Reserve,Gabon

Sympatric populations of lowland gorillas (Gorilla gorilla gorilla) and chimpanzees (Pan troglodytes troglodytes) in the Lopé Reserve in central Gabon consumed insects at similar average frequencies over a 7-year period (30% versus 31% feces contained insect remains). Data came mostly from fecal analysis supplemented by observation and trail evidence. The weaver ant (Oecophylla longinoda) was the species eaten most frequently by both gorillas and chimpanzees. Other species of insects wore eaten but there was virtually no overlap: Chimpanzees used tools to eat Apis bees (and their honey) and two large species of ants; gorillas ate three species of small ants. Thus, despite their shared habitat, the esources utilized were not identical as gorillas do not show the tool-use “technology” of chimpanzees. The frequency of insect-eating by both species of ape varied seasonally and between years but in different ways. This variation did not seem to be related to the ratio of fruit to foliage in their diets. Gorillas of all age-classes ate insects at similar rates. Comparisons with insectivory by other populations of gorillas indicate differences exist. Mountain gorillas (Gorilla g. beringei) in the Virunga Volcanoes, Rwanda, consume thousands of invertebrates daily, eating them inadvertently with handfuls of herbaceous foods but they deliberately ingest insect-foods only rarely. Lowland gorillas at Lopé habitually ate social insects, and their selective processing of herbaceous foods probably minimizes inadvertent consumption of other invertebrates. Gorillas at Belinga in northeastern Gabon, 250 km from Lop6, ate social insects at similar rates but ignored weaver ants in favor of Cubitermes sulcifrons, a small species of termite that occurs at Lopé but was not eaten by gorillas. This indicates that local traditions similar to those reported for chimpanzees also exist amongst populations of gorillas. © 1992 Wiley-Liss, Inc.     

Leaf Surface Roughness Elicits Leaf Swallowing Behavior in Captive Chimpanzees (Pan troglodytes) and Bonobos (P. paniscus), but not in Gorillas (Gorilla gorilla) or Orangutans (Pongo abelii)

Researchers have described apparently self-medicative behaviors for a variety of nonhuman species including birds and primates. Wild chimpanzees, bonobos, and gorillas have been observed to swallow rough leaves without chewing, a behavior proposed to be self-medicative and to aid control of intestinal parasites. Researchers have hypothesized that the presence of hairs on the leaf surface elicits the behavior. We investigated the acquisition and the underlying mechanisms of leaf swallowing. We provided 42 captive great apes (24 chimpanzees, six bonobos, six gorillas, and six orangutans) with both rough-surfaced and hairless plants. None of the subjects had previously been observed to engage in leaf swallowing behavior and were therefore assumed naïve. Two chimpanzees and one bonobo swallowed rough-surfaced leaves spontaneously without chewing them. In a social setup six more chimpanzees acquired the behavior. None of the gorillas or orangutans showed leaf swallowing. Because this behavior occurred in naïve individuals, we conclude that it is part of the behavioral repertoire of chimpanzees and bonobos. Social learning is thus not strictly required for the acquisition of leaf swallowing, but it may still facilitate its expression. The fact that apes always chewed leaves of hairless control plants before swallowing, i.e., normal feeding behavior, indicates that the surface structure of leaves is indeed a determinant for initiating leaf swallowing in apes where it occurs.     

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. Am. J. Primatol. 71:91–100, 2009. © 2008 Wiley‐Liss, Inc.     

Use of space in a non-naturalistic environment by chimpanzees (Pan troglodytes) and lowland gorillas (Gorilla gorilla gorilla)

Chimpanzees and gorillas are the two most common species of great ape in captive facilities in North America. This study examined patterns of space use by 14 gorillas and six chimpanzees housed in similar non-naturalistic environments at Lincoln Park Zoo in Chicago, IL. The location of each individual was recorded in relation to elements of the environment over a two-year period. These data were compared to volumetric measurements of the enclosures to determine “preferences” for particular environmental elements. Chimpanzees preferred the highest tier of the enclosure and the gorillas preferred the floor level. Both species showed preferences for doorways, corners and the mesh barriers adjacent to keeper areas. These data supplement data from wild populations of apes and provide information useful for those seeking to design new ape enclosures or renovate existing facilities.     

Dehydroepiandrosterone-sulfate (DHEA-S), sex,and age in zoo-housed western lowland gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>)

Among humans, dehydroepiandrosterone-sulfate (DHEA-S) declines with age and is hypothesized to be involved in somatic maintenance and healthy aging. Men have significantly higher DHEA-S than women, contradicting longer lifespans in the latter. Declines of DHEA-S with age also are observed in chimpanzees. In both chimpanzees and bonobos, males and females show no differences in DHEA-S production. Based on human and chimpanzee data, gorillas were predicted to show declining DHEA-S with age. Similar to chimpanzees and bonobos, it also was predicted DHEA-S would not be significantly different between males and females. DHEA-S was assayed from serum banked during physical examinations of gorillas housed at three North American zoos (n = 63). Gorillas ranged from 6 to 52 years of age. Differences between males and females were examined using t tests. Linear regression was used to determine the relationship of DHEA-S with age. There was no significant difference in DHEA-S between males and females. Additionally, there was no significant relationship between DHEA-S and age. As predicted, there were no sex-based differences in DHEA-S in gorillas, which is similar to chimpanzees and bonobos but different from modern humans. Unlike chimpanzees and humans, there was no significant relationship between DHEA-S and age in gorillas. The absence of a relationship between age and DHEA-S may be due to the lack of gorillas under age 6 years in this sample as declines in chimpanzees occur prior to age 5 years, more rapid growth and development among gorillas compared with other African hominoids, or a unique pattern of DHEA-S production.     

Brain structure variation in great apes, with attention to the mountain gorilla (Gorilla beringei beringei)

This report presents data regarding the brain structure of mountain gorillas (Gorilla beringei beringei) in comparison with other great apes. Magnetic resonance (MR) images of three mountain gorilla brains were obtained with a 3T scanner, and the volume of major neuroanatomical structures (neocortical gray matter, hippocampus, thalamus, striatum, and cerebellum) was measured. These data were included with our existing database that includes 23 chimpanzees, three western lowland gorillas, and six orangutans. We defined a multidimensional space by calculating the principal components (PCs) from the correlation matrix of brain structure fractions in the well-represented sample of chimpanzees. We then plotted data from all of the taxa in this space to examine phyletic variation in neural organization. Most of the variance in mountain gorillas, as well as other great apes, was contained within the chimpanzee range along the first two PCs, which accounted for 61.73% of the total variance. Thus, the majority of interspecific variation in brain structure observed among these ape taxa was no greater than the within-species variation seen in chimpanzees. The loadings on PCs indicated that the brain structure of great apes differs among taxa mostly in the relative sizes of the striatum, cerebellum, and hippocampus. These findings suggest possible functional differences among taxa in terms of neural adaptations for ecological and locomotor capacities. Importantly, these results fill a critical gap in current knowledge regarding great ape neuroanatomical diversity.     

A Survey of Chimpanzees (Pan troglodytes troglodytes) and Gorillas (Gorilla gorilla gorilla) in the Selectively Logged Ngotto Forest,Central African Republic

Currently, the timber company Industrie Forestiére du Batalimo is selectively logging the Ngotto Forest in the Central African Republic. The forest is home to a population of chimpanzees (Pan troglodytes troglodytes) and gorillas (Gorilla gorilla gorilla) and provides the Bofi-speaking people of Grima with food, medicine, housing material, and other commodities. Over a 7-month period, the research team conducted a line-transect survey of the great ape population in the forests to the south of Grima to document their distribution. For comparison purposes, the team also surveyed a section of adjacent forest that had already been logged. Ape nests were significantly rarer in the logged forest than in the unlogged forest, and ape nests were most common in the more pristine forests to the south. This report further discusses the effects of logging and other human activities.     

Species association of hepatitis B virus (HBV) in non-human apes; evidence for recombination between gorilla and chimpanzee variants

Hepatitis B virus (HBV) infections are widely distributed in humans, infecting approximately one third of the world's population. HBV variants have also been detected and genetically characterised from Old World apes; Gorilla gorilla (gorilla), Pan troglodytes (chimpanzee), Pongo pygmaeus (orang-utan), Nomascus nastusus and Hylobates pileatus (gibbons) and from the New World monkey, Lagothrix lagotricha (woolly monkey). To investigate species-specificity and potential for cross species transmission of HBV between sympatric species of apes (such as gorillas and chimpanzees in Central Africa) or between humans and chimpanzees or gorillas, variants of HBV infecting captive wild-born non-human primates were genetically characterised. 9 of 62 chimpanzees (11.3%) and two from 11 gorillas (18%) were HBV-infected (15% combined frequency), while other Old world monkey species were negative. Complete genome sequences were obtained from six of the infected chimpanzee and both gorillas; those from P. t .ellioti grouped with previously characterised variants from this subspecies. However, variants recovered from P. t. troglodytes HBV variants also grouped within this clade, indicative of transmission between sub-species, forming a paraphyletic clade. The two gorilla viruses were phylogenetically distinct from chimpanzee and human variants although one showed evidence for a recombination event with a P.t.e.-derived HBV variant in the partial X and core gene region. Both of these observations provide evidence for circulation of HBV between different species and sub-species of non-human primates, a conclusion that differs from the hypothesis if of strict host specificity of HBV genotypes.     

Another gorilla (Gorilla gorilla gorilla) recognizes himself in a mirror

Various attempts have been made to explain why gorillas (Gorilla gorilla gorilla) find it difficult to recognize their mirror image. One of the most oft-cited reasons is aversion to eye contact, which stops gorillas from looking into a mirror and thus prevents them from carrying out a suitable exploration that could lead to self-recognition. In the experimental design used here the subject was first habituated both to observers and to the mirror as an object before being exposed to the latter. The study was performed with a single subject who was well adapted to captivity and exhibited no aberrant behavior or signs of stress. The results revealed that the subject had no aversion to eye contact. He showed considerable interest in the mirror and appeared relaxed when faced with his image. He gave a positive response to the mark test.     

ORIGINAL ARTICLE: Medical and surgical management of reproductive neoplasia in two western lowland gorillas (Gorilla gorilla gorilla)

Background Reports of female reproductive tract neoplasia are infrequent in great apes. Methods Two captive‐born, female western lowland gorillas (Gorilla gorilla gorilla) housed at the North Carolina Zoological Park were diagnosed with reproductive neoplasia. Results The first gorilla had uterine endometrial thickening and uterine fibroids diagnosed during a routine annual examination. Subsequently, the animal underwent several uterine curettage procedures in an attempt to debulk the uterine mass. Biopsy results indicated uterine adenocarcinoma. The animal was treated with an ovariohysterectomy followed by a combination of megestrol acetate and medroxyprogesterone acetate. The second gorilla was diagnosed with squamous cell carcinoma of the vagina, cervix, and uterus after presenting for intermittent vaginal bleeding of 1‐month duration. This animal underwent palliative radiation therapy, receiving two rounds of radiation treatment 8 weeks apart. Conclusions The following report describes the diagnosis and management of the two cases.     

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.     

Sexual behavior in female western lowland gorillas (Gorilla gorilla gorilla): evidence for sexual competition

Previous research in gorillas suggests that females engage in post‐conception mating as a form of sexual competition designed to improve their own reproductive success. This study focused on sexual behaviors in a newly formed group of western lowland gorillas (Gorilla gorilla gorilla) housed at Zoo Atlanta. All females engaged in mating outside their conceptive periods, although there was individual variation in the frequency of the behavior. An analysis of the presence/absence of sexual behavior found females, regardless of reproductive condition, were more likely to engage in sexual behavior on days when other females were sexually active. On these “co‐occurrence” days, females were significantly more likely to solicit the silverback, but copulations did not differ from expectation. The results find further evidence for sexual competition among female gorillas and suggest that this may occur throughout their reproductive cycle rather than only during pregnancy. Am. J. Primatol. 71:587–593, 2009. © 2009 Wiley‐Liss, Inc.     

Hand Preference for a Novel Bimanual Coordinated Task During Termite Feeding in Wild Western Gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>)

Although the level of handedness in humans varies cross-culturally, humans are generally described as right-handed, which has been considered a uniquely human trait. Recently, captive chimpanzees (Pan troglodytes) have been shown to exhibit right-hand preference when performing bimanual but not unimanual tasks. Less clear is whether this pattern also occurs in wild chimpanzees and other African apes. Using videos (N = 49) of six wild western gorillas (Gorilla gorilla gorilla) feeding on termites at the Mondika Research Center (Republic of Congo), we tested whether they exhibit hand preference when performing unimanual, i.e., reaching for termite mound pieces; bimanual, i.e., “termite tapping”: rhythmically shaking a piece of termite mound with the dominant hand and collecting the termites in the other hand tasks; or hand transfer prior to bimanual tasks, i.e., transferring a piece of termite mound from one hand to the other. All individuals exhibited exclusive hand preference when performing the bimanual tasks, with five of six gorillas preferring the right hand. Conversely, most individuals did not show any manual preference during the unimanual task. In addition, hand preference during hand transfer revealed clear hand dominance of similar strength and direction of those shown for the bimanual task, suggesting that this measure is as sensitive as the bimanual task itself. Thus, we propose “termite feeding” as a novel task to be considered in future hand-preference studies in wild western gorillas. Our results are in concordance with those for chimpanzees and captive gorillas showing hemispheric specialization for bimanual actions in apes.     

Gestural communication in young gorillas (Gorilla gorilla): gestural repertoire,learning, and use

In the present study we investigated the gestural communication of gorillas (Gorilla gorilla). The subjects were 13 gorillas (1-6 years old) living in two different groups in captivity. Our goal was to compile the gestural repertoire of subadult gorillas, with a special focus on processes of social cognition, including attention to individual and developmental variability, group variability, and flexibility of use. Thirty-three different gestures (six auditory, 11 tactile, and 16 visual gestures) were recorded. We found idiosyncratic gestures, individual differences, and similar degrees of concordance between and within groups, as well as some group-specific gestures. These results provide evidence that ontogenetic ritualization is the main learning process involved, but some form of social learning may also be responsible for the acquisition of special gestures. The present study establishes that gorillas have a multifaceted gestural repertoire, characterized by a great deal of flexibility with accommodations to various communicative circumstances, including the attentional state of the recipient. The possibility of assigning Seyfarth and Cheney's [1997] model for nonhuman primate vocal development to the development of nonhuman primate gestural communication is discussed.     

Sex-biased dispersal in western lowland gorillas (Gorilla gorilla gorilla)

We explored two hypotheses related to potential differences between sexes in dispersal behaviour in western lowland gorillas (Gorilla gorilla gorilla). Direct observations suggest that immature females have more opportunities to move between breeding groups than immature males. The distribution of kin dyadic relationships within and between groups does not, however, support this hypothesis. At larger geographical scales, dispersal is likely to be easier for males than females because of the solitary phase most blackbacks experience before founding their own breeding group. However, previous work indicates that males settle preferentially close to male kin. By specifically tracing female and male lineages with mitochondrial and Y-chromosomal genetic markers, we found that male gorillas in the 6000 km2 area we surveyed form a single population whereas females are restricted to the individual sites we sampled and do not freely move around this area. These differences are more correctly described as differences in dispersal distances, rather than differences in dispersal rates between sexes (both sexes emigrate from their natal group in this species). Differences in resource competition and dispersal costs between female and male gorillas are compatible with the observed pattern, but more work is needed to understand if these ultimate causes are responsible for sex-biased dispersal distances in western lowland gorillas.