Newly Discovered Gorilla Population in the Ebo Forest,Littoral Province,Cameroon

We report on a new population of gorillas discovered in November 2002 in the Ebo Forest, Littoral Province, Cameroon. We observed A group of q7 gorillas directly for 83 min, and they were in auditory range for 155 min. Further evidence of gorilla presence included 8 nest groups totaling 38 nests, distinctive feeding signs accompanied by footprints, and a gorilla cranium collected from the nearby village of Iboti. This newly discovered gorilla population is geographically intermediate between the 2 extant populations of western gorillas: Gorilla gorilla gorilla, the most populous gorilla subspecies living in Gabon, Equatorial Guinea, Congo-Brazzaville, Central African Republic and Cameroon to the south of the Sanaga River, and G. g. diehli or the Cross River gorilla, a small population of ca. 250 individuals on the Cameroon-Nigeria border. It is not possible to assign the new gorilla population to either subspecies on the basis of measurements of the single male cranium. Genetic analyses of freshly shed hairs, collected from gorilla nests, may help to resolve the taxonomic status of the Ebo gorillas.     

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.     

Termite Feeding by <Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis> at Bai Hokou,Central African Republic

Though insectivory by large-bodied gorillas may be unexpected, researchers have reported it in all populations of gorillas studied to date. Our study of 2 well monitored groups of western gorillas (Gorilla gorilla gorilla) at Bai Hokou in Dzanga-Ndoki National Park, Central African Republic provides information on frequency and variability of termite consumption (the most commonly eaten insect) as well as some of the first direct observations of the behavior. Pooled data from both groups indicate termite feeding on 34% and 83% of days, through fecal analysis and feeding trails, respectively. Direct observations revealed that termite feeding occurred on 91% of the days for 1 group, in which the silverback fed on termites during 13% of all feeding scans, making termites the most commonly observed food item. The group that had a higher density of termite mounds in its home range consumed termites more frequently than the other group did. A higher proportion of fecal samples from the silverbacks contained termite remains than the ones from adult females and juveniles. Termite consumption was lower during the dry season, but it does not correlate with rainfall, measures of fruit availability, or fruit consumption. Displacements at termite mounds occurred more than expected, indicating that they are a patchy, sought-after food resource. Gorillas did not use tools to extract termites, but they used 2 different techniques to remove them from the cells. Though culture or social traditions may cause the variation in termite consumption across sites, further investigation of termite availability and consumption is necessary to rule out ecological and methodological explanations for observed variations.     

Gorillas (Gorilla gorilla gorilla) in the Likouala swamp forests of north central Congo: Preliminary data on populations and ecology

Until recently it was generally believed that gorillas avoid water. In two recent, independent faunal surveys in the Likouala swamp of north central Congo, we have found that gorillas occur at high densities, extending the known range of Gorilla gorilla gorillavery close to the Oubangui River. The gorillas in this area fed on the abundant ground vegetation of herbaceous monocotyledons in the swamp forest. In adjacent islands of terra firmathere was little evidence of gorilla nesting or feeding, indicating that they spend at least part of the year almost exclusively in the swamp forests. Recent reports from several sites in West Africa indicate that swamp forests may generally provide abundant food resources that are used regularly by gorillas.     

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.     

Mitochondrial DNA sequence from an enigmatic gorilla population (Gorilla gorilla uellensis)

Although today gorillas are found in only two widely separate, discontinuous western and eastern African populations, rumors of the existence of an additional gorilla population in central Africa have inspired recent unsuccessful field expeditions in search of the "mystery ape" termed Gorilla gorilla uellensis. Such a gorilla population would have considerable conservation and scientific interest, and would presumably have descended from a population of gorillas that was thought to exist until the end of the 19th century on the Uele River in the current-day Democratic Republic of Congo. However, the sole evidence for the existence of these gorillas is three skulls and one mandible brought to the Royal Museum for Central Africa (Tervuren, Belgium) in 1898. We determined a mitochondrial DNA sequence from one of these specimens and compared it to sequences from other gorillas. Contrary to expectations, the sequence obtained did not exhibit the phylogenetic distinctiveness typical of a representative of a peripheral isolated population. Rather, the results suggest a scenario in which the museum specimens did not originally derive from the northern Congo, but were brought from the area of current distribution of western gorillas to that location; the subsequent discovery and collection of the specimens there gave rise to the false inference of a local gorilla population.     

Taste Responses to Fructose and Tannic Acid Among Gorillas (Gorilla gorilla gorilla)

We investigated the taste responses to fructose and tannic acid compounds among 6 western gorillas (Gorilla gorilla gorilla) at the San Francisco Zoo. We presented the subjects with 12 reference concentrations of 3–300 mM fructose, paired with tap water in paired-choice experimental trials. We subsequently presented them with tap water paired with 8 dilute concentrations of tannic acid (0.25–6 mM) dissolved in 100 mM fructose solutions and 8 concentrations (0.25–6 mM) dissolved in 300 mM fructose solutions. The gorillas exhibited a broadly similar preference threshold for fructose (50 mM) to those of other nonhuman primate and human samples. The gorillas tolerated moderate levels of tanninc acid, especially when presented in a very sweet package. The depressing effect of tannic acid on the ingestion of fructose solutions appeared to increase progressively with tannin concentration and was lower as fructose concentration increased. The inhibition threshold for tannic acid solutions was reached at 4 mM tannic acid in 100 mM fructose solution, and at 6 mM tannic acid in 300 mM fructose solutions. These results suggest that gorillas use sweetness as a criterion for food selection and regard those with moderate levels of tannins as palatable. Our findings corroberate food preference studies and nutritional analyses of wild gorilla foods indicating that they prefer sugary foods and readily consume ones containing moderate levels of tannins. Taste responses may facilitate a flexible frugivorous/folivorous diet among gorillas.     

Ant eating behavior of mountain gorillas

Eleven cases of feeding on driver ants (Dorylus sp.) by mountain gorillas (Gorilla gorilla beringei) are described. Ant eating provides the gorillas with more animal protein and other nutrients per unit feeding time than do other forms of insectivory that contribute to their diet, but it is so rare that it is unlikely to be of real nutritional significance. Gorillas obtain ants with their hands and do not use tools. Immature individuals (except infants) ate more ants than did adult females, and silverbacks were not seen to eat ants. These differences are more likely to reflect differences in individual taste and interest in novelty than differences in nutritional strategy. Not all gorillas in the Virungas population eat ants. Intra-population variability may be ecologically contingent, but ant eating appears to be a socially acquired and transmitted taste.     

Seasonality in the Ecology and Life Histories of Mountain Gorillas (Gorilla gorilla beringei)

The abundance of food, especially that of fruit and often that of young leaves, varies considerably over time for most primates. This variation can depend on or be independent of seasonality in rainfall. Mountain gorillas (Gorilla gorilla beringei) in the Virungas are exceptional: their habitat contains almost no edible fruit, and they mostly eat perennially available herbs and vines that are densely and evenly distributed in much of their habitat. Earlier studies documented little consistent temporal variation in mountain gorilla diets and habitat use, except for seasonal use of bamboo by some groups, and documented no birth seasonality. Long-term data ( 7 years) on 6 mountain gorilla groups confirm these results for habitat use, except that they show some seasonality in use of the upper altitudinal extremes of the gorillas' home ranges for unclear reasons. Relatively low and inconsistent variation in habitat quality over time should lower the costs of grouping for gorillas compared to other apes. Long-term data also confirm the absence of seasonality in births and conceptions. However, they show that mortality rates and risk of respiratory infections vary directly with rainfall. These relationships are probably causal and may be mediated through thermoregulatory stress.     

Monitoring the reproductive status of female gorillas (Gorilla gorilla gorilla) by measuring the steroid hormones in fecal samples

The reproductive status of female gorillas (Gorilla gorilla gorilla) was estimated by measuring the sex steroid hormones in fecal samples instead of in blood samples. Fecal samples from female gorillas were used to examine the reliability of this non-invasive assay system, which included the extraction method for estradiol-17β (E₂) and progesterone (P) from fecal samples. Fecal samples from three female gorillas were collected daily for about 55 days, and fecal E₂ and P were assayed to clarify the fluctuation patterns of these steroids in the feces. Fecal sampling from one female was repeated for another 50-day period (starting 75 days after the end of the first observation period) and assayed to confirm if the menstrual cycle of this subject was ovulatory. Although fecal E₂ concentration measurements were quantitative by using this assay system, fecal P concentration measurements were semi-quantitative. Relative amounts of fecal P in fecal samples were estimated by using the values of B/B₀ (bound/total binding in the radioimmunoassay system). Two of the four fluctuation patterns of fecal hormones observed throughout the menstrual cycle for the three female gorillas were typical for normal ovulatory cycles. In the subject observed for two periods, one pattern was typical and the other atypical. The results show that this non-invasive method is simple and practical for monitoring the reproductive status of great apes as well as Old World monkeys.     

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.     

Mitochondrial COII Introgression into the Nuclear Genome of Gorilla gorilla

Numts are nonfunctional mitochondrial sequences that have translocated into nuclear DNA, where they evolve independently from the original mitochondrial DNA (mtDNA) sequence. Numts can be unintentionally amplified in addition to authentic mtDNA, complicating both the analysis and interpretation of mtDNA-based studies. Amplification of numts creates particular issues for studies on the noncoding, hypervariable 1 mtDNA region of gorillas. We provide data on putative numt sequences of the coding mitochondrial gene cytochrome oxidase subunit II (COII). Via polymerase chain reaction (PCR) and cloning, we obtained COII sequences for gorilla, orangutan, and human high-quality DNA and also from a gorilla fecal DNA sample. Both gorilla and orangutan samples yielded putative numt sequences. Phylogenetically more anciently transferred numts were amplified with a greater incidence from the gorilla fecal DNA sample than from the high-quality gorilla sample. Data on phylogenetically more recently transferred numts are equivocal. We further demonstrate the need for additional investigations into the use of mtDNA markers for noninvasively collected samples from gorillas and other primates.     

Environmental influences on mountain gorilla time budgets

Data on the time budgets of mountain gorillas (Gorilla gorilla beringei) were collected during field studies in the Virunga Volcanoes region of Rwanda and Zaire. Focal sampling was used to determine the proportion of time that individuals of different age/sex classes spent in several mutually exclusive activity states. The gorillas spent the majority of daylight hours feeding; most of the rest of the day was devoted to resting, with little time spent moving or engaged in social activity. The time budget varied among the different subhabitats used by the gorillas, and the gorillas satisfied subsistence needs more quickly when in areas where food was more abundant and/or of better nutritional quality. Silverbacks spent more time feeding than all other age/sex classes, but age/sex class differences were not great. All age/sex classes responded to variability in habitat quality in similar fashion. Unlike the case for many other primates, there was no significant seasonal variation in time budgets. There was a direct relationship between group size and time spent feeding, although variation in relation to group size was lower than that in relation to variation in habitat quality. These results are consistent with the relationship of feeding time to body size in primates. They are also consistent, with other evidence that social foraging entails a cost to gorilla females, but that this cost is low in comparison to those faced by many other primates. Permanent association with males apparently offers little ecological disadvantage to females, who are likely to be more than compensated by mutualistic benefits.     

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.     

Behavioral and hormonal responses to the availability of forage material in Western lowland gorillas (Gorilla gorilla gorilla)

We investigated how forage material affects indicators of welfare in three male Western lowland gorillas (Gorilla gorilla gorilla) at the Detroit Zoo. In addition to their maintenance diet and enrichment foods, the gorillas generally received forage material four times a week. From this baseline, we systematically manipulated how much forage material the group received on a weekly basis, with either daily or bi (twice)‐weekly presentation of browse (mulberry, Morus sp.) or alfalfa hay. We collected behavioral data (60 hr per gorilla) and measured fecal glucocorticoid metabolites (FGM). Mixed models indicated that the presence of forage material significantly increased time feeding (F_2,351 = 9.58, p < 0.001), and decreased rates of noncontact aggression (F_2,351 = 3.69, p = 0.03), and regurgitation and reingestion (F_2,353 = 4.70, p = 0.01). Regurgitation and reingestion were never observed during the condition when forage material was provided daily. When forage material was provided, time spent feeding was similar across gorillas, compared to a disproportionately greater amount of time spent feeding by the dominant individual when forage material was absent. Providing forage material in addition to the regular diet likely created more opportunities for equitable feeding for the subordinate gorillas. FGM concentrations did not vary based on the presence or type of forage material available and, instead, likely reflected group social dynamics. In general, alfalfa and mulberry had similar impacts on behavior, indicating that alfalfa can be an adequate behavioral substitute during times when browse is less readily available for gorillas housed in seasonally variable climates.     

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.     

Male and female western gorilla diet: Preferred foods,use of fallback resources,and implications for ape versus old world monkey foraging strategies

Most of what is currently known about western gorilla (Gorilla gorilla) diet is based on indirect studies using fecal samples and trail signs rather than measures based on direct observations. Here we report results on adult male and female western gorilla foraging behavior, based on systematic focal observations and nutritional analyses of foods. We found that western gorillas, like other apes, are highly selective ripe fruit specialists, seeking fruit high in energy, low in antifeedants, and rare in the environment. During seasonal fruiting peaks, fruit accounted for up to 70% of feeding time. When ripe fruit was scarce, gorillas increased time spent feeding on leaves and nonpreferred fruits and herbs. Leaves were the major fallback food, accounting for up to 70% of feeding time in males and 50% in females during periods of fruit scarcity. In spite of large differences in body size, the sexes were remarkably similar in their overall diet, not differing in time spent feeding on fruit or preferred herbs. However, the male consistently fed more often and on a greater variety of leaves than did females, whereas females fed more often on fallback herbs and termites. Our findings, when considered in light of previous findings on sympatric mangabeys, indicate that the foraging strategy of western gorillas is broadly similar to that of chimpanzees and orangutans, and distinct from that of old world monkeys. Am J Phys Anthropol 140:727–738, 2009. © 2009 Wiley-Liss, Inc.     

The Intestinal Entodiniomorph Ciliates of Wild Lowland Gorillas (Gorilla gorilla gorilla) in Gabon,West Africa

Examination of 109 fecal samples from wild lowland gorillas revealed the presence of five species of entodiniomorph ciliates: Troglodytella ahrassarti, Troglodytella gorillae, and three unclassified species. These latter three species were also found in the feces of a captive gorilla in Gabon and are considered to be intestinal parasites or commensals.     

The cryptic genetic structure of the North American captive gorilla population

Western lowland gorillas (Gorilla gorilla gorilla) were imported from across their geographical range to North American zoos from the late 1800s through 1974. The majority of these gorillas were imported with little or no information regarding their original provenance and no information on their genetic relatedness. Here, we analyze 32 microsatellite loci in 144 individuals using a Bayesian clustering method to delineate clusters of individuals among a sample of founders of the captive North American zoo gorilla collection. We infer that the majority of North American zoo founders sampled are distributed into two distinct clusters, and that some individuals are of admixed ancestry. This new information regarding the existence of ancestral genetic population structure in the North American zoo population lays the groundwork for enhanced efforts to conserve the evolutionary units of the western lowland gorilla gene pool. Our data also show that the genetic diversity estimates in the founder population were comparable to those in wild gorilla populations (Mondika and Cross River), and that pairwise relatedness among the founders is no different from that expected for a random mating population. However, the relatively high level of relatedness (R = 0.54) we discovered in a pair of known breeding pairs reveals the need for incorporating genetic relatedness estimates in the captive management of western lowland gorillas.     

Species and sex identification of western lowland gorillas (Gorilla gorilla gorilla), eastern lowland gorillas (Gorilla beringei graueri) and humans

Methods for the identification of the sex and species of individuals from samples non-invasively taken from humans and gorillas were established. Amplification of a segment of amelogenin (AMG), which is an X–Y homologous gene, using two pairs of primers from human AMG, revealed both X- and Y-specific bands. The possibility of sex identification was examined by typing the AMG gene using hair and fecal samples from captive western lowland gorillas (Gorilla gorilla gorilla) in Japan and hair samples from wild eastern lowland gorillas (Gorilla beringei graueri) in the Kahuzi-Biega National Park, Democratic Republic of Congo, which were sexed by direct observation. Species-specific bands of AMG in gorillas and humans were identified by restriction fragment length polymorphisms analysis. These tests could be used for sexing unidentified individuals of wild western and eastern lowland gorillas, and screening contamination of human DNA from non-invasively acquired samples.