Are the gorillas in Bwindi Impenetrable National Park “true” mountain gorillas?

The gorillas that inhabit Bwindi Impenetrable National Park in Uganda are the least known of the eastern gorillas. Because they are an allopatric population living a minimum of 25 km from the well‐studied population of mountain gorillas (Gorilla beringei beringei) in Rwanda and have certain morphological and ecological differences from these gorillas, their taxonomic status has been in question in recent years. This study presents new craniodental metrics from Bwindi individuals and compares them to Virunga individuals as well as to eastern lowland gorillas, G. gorilla graueri. Multivariate statistics, including MANCOVA, least‐squares, regression, and principal components analyses, were used to evaluate how closely the Bwindi crania resemble the Virunga crania and how both relate to G. g. graueri. Results indicate that the Bwindi gorillas have generally smaller crania than the Virunga gorillas, but when metrics are log‐transformed, the only variable that distinguishes the Bwindi individuals is a longer face. When both populations are compared to G. g. graueri, they cluster together separately from the eastern lowland gorillas, sharing such features as higher rami, wider bigonia, longer mandibles, and wider and shorter mandibular symphyses in relation to G. g. graueri. Functional morphological explanations for these differences are discussed, but lacking measurements of the physical properties of G. g. graueri, they cannot fully be explained. Results clearly indicate that at least pertaining to the cranium, upon which most gorilla taxonomy is based, the Bwindi gorillas are proper mountain gorillas (G. b. beringei). Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

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

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

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.     

Home Range and Frugivory Patterns of Mountain Gorillas in Bwindi Impenetrable National Park,Uganda

Mountain, western, and Grauer's gorillas exhibit broad differences in ecological patterns with western gorillas eating more fruit and having larger home ranges than their largely folivorous counterparts in the Virunga Volcanoes. We studied the home range and frugivory patterns of one group of Gorilla beringei beringei in the little-studied population of Bwindi Impenetrable National Park, Uganda, to compare with other populations and to investigate whether there was any relationship between patterns of frugivory and home range size. During the 3-year study, the gorillas ate 16 species of fruit on 27% of observation days. There was high variability in frugivory among the 3 years and no consistent seasonal pattern. Annual home range size was ca. 21 km² for Years 1 and 2, and it increased dramatically to 40 km² in Year 3. Home range size varied considerable between months and seasons, but there is no clear relationship between occurrence of fruit-eating and home range size. The group exhibited more fruit-eating and a larger home range size those ofthe gorillas in the Virunga Volcanoes. Their home range size is comparable to that of western gorillas, though Bwindi gorillas consumed less fruit. Home range size and utilization by all gorillas probably depends on a complex relationship between the distribution and abundance of both fruit and herbaceous vegetation and social factors such as male mating tactics.     

Mitochondrial DNA phylogeography of western lowland gorillas (Gorilla gorilla gorilla)

The geographical distribution of genetic variation within western lowland gorillas (Gorilla gorilla gorilla) was examined to clarify the population genetic structure and recent evolutionary history of this group. DNA was amplified from shed hair collected from sites across the range of the three traditionally recognized gorilla subspecies: western lowland (G. g. gorilla), eastern lowland (G. g. graueri) and mountain (G. g. beringei) gorillas. Nucleotide sequence variation was examined in the first hypervariable domain of the mitochondrial control region and was much higher in western lowland gorillas than in either of the other two subspecies. In addition to recapitulating the major evolutionary split between eastern and western lowland gorillas, phylogenetic analysis indicates a phylogeographical division within western lowland gorillas, one haplogroup comprising gorilla populations from eastern Nigeria through to southeast Cameroon and a second comprising all other western lowland gorillas. Within this second haplogroup, haplotypes appear to be partitioned geographically into three subgroups: (i) Equatorial Guinea, (ii) Central African Republic, and (iii) Gabon and adjacent Congo. There is also evidence of limited haplotype admixture in northeastern Gabon and southeast Cameroon. The phylogeographical patterns are broadly consistent with those predicted by current Pleistocene refuge hypotheses for the region and suggest that historical events have played an important role in shaping the population structure of this subspecies.     

Movement patterns of a group of free-ranging mountain gorillas (Gorilla gorilla beringei) and their relation to food availability

Relationships between the movement patterns of free-ranging mountain gorillas (Gorilla gorilla beringei) and the abundance and distribution of foods in their home range were examined. During an 18-month field study, the ranging of one group of G. gorilla beringei was recorded on a 250 × 250 m grid system, from which measurements of frequency and duration of use, travel rate, and rate of revisitation of each quadrangle by the group were derived. Food items were sampled in selected quadrangles throughout the home range and various measures of food abundance, frequency and diversity were calculated. Analyses based on both spatial and temporal variation in food availability give supporting evidence for the prediction that mountain gorilla ranging patterns are influenced by the distribution and abundance of foods. Quality of food appears to be an important factor, as shown in analyses of yearly patterns and monthly changes in ranging. The gorillas studied spent more time in the higher quality areas of their home range, responded to a correlate of decreasing food abundance by increasing their rate of travel and area used, and revisited regions more frequently when the renewal rate of foods was clearly greater. Each of their foraging tactics can be explained as serving to increase the efficiency of harvesting foods.     

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.     

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.     

Long‐term proximity relationships in a captive social group of western lowland gorillas (Gorilla gorilla gorilla)

To visualize long‐term social relationships among 12 gorillas in a captive breeding group at the San Diego Wild Animal Park, San Diego, CA, the multidimensional scaling (MDS) procedure was applied to proximity (within 5 m) values collected in five different periods over 2.5 years. The resulting two‐dimensional representations clearly show that in samples taken during this 2.5‐year period, the silverback male was surrounded by five adult females, while three infant or young juvenile females, gathered in a body, remained between the silverback male and adult females. Some adult females maintained proximity to the silverback male more frequently than other adult females throughout the five periods. Unlike mountain gorillas in the wild, females with dependent offspring did not tend to stay near the silverback male more frequently than other females, and related females did not tend to spend more time near each other than non‐related females. Three older juvenile or young adult males were plotted the furthest from the silverback male, with gradual changes with increasing age. The usefulness of the MDS procedure, which can be used to visualize easily and clearly the social relationships among individuals, is discussed from the viewpoint of the management of breeding groups of captive gorillas. Zoo Biol 20:197–209, 2001. © 2001 Wiley‐Liss, Inc.     

Inter-group encounters and female transfer in mountain gorillas: Influence of group composition on male behavior

This paper examines the behaviors used by male gorillas (G. gorilla beringei) to influence female choice during inter-group encounters. Encounters are related to the acquisition of females rather than to the defense of a group's range. Data on 58 independent encounters confirms that encounters are generally aggressive, but that contact aggression is less frequent than previously reported. A high number of potential migrants in one of the groups involved, rather than the newness of either group, predicts the likelihood of contact aggression between males. Encounters with a newly formed unit last longer than encounters involving established groups. Male herding serves to prevent female transfer. Females without dependent offspring are more likely than others to be herded. Proceptive females are the target of herding more often than non-proceptive cycling females. Groups where herding has been observed are more likely to be newly formed, and they include significantly more males. This last result confirms that there are advantages to male cooperation in gorillas. It also raises the question of why such cooperation does not occur more often. © 1993 Wiley-Liss, Inc.     

Ranging and grouping patterns of a western lowland gorilla group at Bai Hokou,Central African Republic

The ranging and grouping patterns of a gorilla group were studied during 27 months from 1990–1992 at the Bai Hokou study site, Central African Republic. The study group ranged far daily (average = 2.3 km/day) and had a large home range (22.9 km²), relative to mountain gorillas, and ranging patterns differed between years. During 1990–1992, the bimale study group foraged less cohesively and had more flexible grouping patterns than mountain gorillas. The study group sometimes split into two distinct foraging subgroups, each led by a silverback, and these subgroups occasionally slept apart (mean = 950 m apart). Lowland gorillas rely on many of the same fruit resources as sympatric chimpanzees, and under certain demographic situations gorillas, like sympatric chimpanzees, may adapt their foraging group size to reduce intragroup feeding competition. However, the fiber content of the lowland gorilla diet likely relaxes constraints on foraging party size and facilitates group cohesion relative to chimpanzees. Am. J. Primatol. 43:111–133, 1997. © 1997 Wiley-Liss, Inc.     

Nutritional chemistry of foods eaten by gorillas in Bwindi Impenetrable National Park, Uganda

Foods eaten by gorillas (Gorilla beringei) in Bwindi Impenetrable National Park (BINP), Uganda, were analyzed for their nutrient content. The goal of the study was to assess the amounts of fiber, protein, and sugars in the foods eaten by the Bwindi gorillas, and to determine whether condensed tannins and cyanide are present in these foods. A total of 127 food plant parts representing 84 plant species eaten by two groups of Bwindi gorillas were collected, processed, and analyzed for their chemical contents. The Bwindi gorilla ate foods that contain 2-28% crude protein (CP), 21-88% neutral detergent fiber (NDF), 14-60% acid detergent fiber (ADF), 2-42% acid detergent lignin (ADL), and 相似文献   

Intrapopulation differences in ant eating in the mountain gorillas of Bwindi Impenetrable National Park, Uganda

Variability in ant eating has been observed in several populations of eastern and western gorillas. We investigated the occurrence of ant (Dorylus sp.) eating in two groups of mountain gorillas (Gorilla beringei beringei) with overlapping home ranges within Bwindi Impenetrable National Park, Uganda from September 2001 to August 2002. We calculated the frequency of ant eating by an indirect method of analyzing fecal samples from silverbacks, adult females, and juveniles. One group consumed ants significantly more often than the other (3.3 vs 17.6% of days sampled). Furthermore, the group that consumed ants more often also consumed them on a seasonal basis (September–February monthly range: 0–8%; March–August monthly range: 30–42.9%). Finally, females and juveniles of this group consumed ants significantly more often than did the silverback (total samples containing ants: silverback, 2.1%; adult female, 13.2%; juvenile, 11.2%). Differences in ant eating between groups are likely due to variability in use of habitats where ants occur (particularly secondary forests). Surveys of ant densities in differing habitats, nutritional analysis of ants, and quantification of the amount of ants in their diets are necessary to understand if ant consumption is due to availability, nutritional value, group traditions, or taste preference.     

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.     

MORPHOLOGICAL AND MOLECULAR EVIDENCE REVEALS RECENT HYBRIDIZATION BETWEEN GORILLA TAXA

Molecular studies have demonstrated a deep lineage split between the two gorilla species, as well as divisions within these taxa; estimates place this divergence in the mid-Pleistocene, with gene flow continuing until approximately 80,000 years ago. Here, we present analyses of skeletal data indicating the presence of substantial recent gene flow among gorillas at all taxonomic levels: between populations, subspecies, and species. Complementary analyses of DNA sequence variation suggest that low-level migration occurred primarily in a westerly-to-easterly direction. In western gorillas, the locations of hybrid phenotypes map closely to expectations based on population refugia and riverine barrier hypotheses, supporting the presence of significant vicariance-driven structuring and occasional admixture within this taxon. In eastern lowland gorillas, the high frequency of hybrid phenotypes is surprising, suggesting that this region represents a zone of introgression between eastern gorillas and migrants from the west, and underscoring the conservation priority of this critically endangered group. These results highlight the complex nature of evolutionary divergence in this genus, indicate that historical gene flow has played a major role in structuring gorilla diversity, and demonstrate that our understanding of the evolutionary processes responsible for shaping biodiversity can benefit immensely from consideration of morphological and molecular data in conjunction.     

Nuclear integrations of mitochondrial DNA in gorillas

Great ape systematics, particularly at the species level and below, is currently under debate, due in part to the recent influx of molecular data. The phylogenies of previously published mitochondrial control region (or D-loop) DNA sequences in gorillas show deep splits within West African gorillas (Gorilla gorilla gorilla), and very high levels of nucleotide diversity in this subspecies. Here we demonstrate that several previously reported D-loop haplotypes from West African gorillas are in all likelihood nuclear integrations of mitochondrial DNA. Revised estimates of the amount and pattern of mitochondrial DNA diversity in gorillas are provided, revealing two reciprocally monophyletic and highly divergent groups of gorillas, concurrent with their geographic distribution.     

Gorillas of Bwindi-Impenetrable Forest and the Virunga Volcanoes: Taxonomic implications of morphological and ecological differences

Based on their geographic proximity to the Virunga Volcanoes (≈ 25 km), the Bwindi-Impenetrable Forest gorillas have been referred to the subspecies Gorilla gorilla beringei. Differences in anatomy, habitat, ecology, and behavior, however, suggest Bwindi gorillas are distinct from those in the Virungas. Relative to Virunga gorillas, Bwindi gorillas live at lower elevations, in warmer temperatures, are much more arboreal, have longer day ranges and larger home ranges, and eat much more fruit and pith, and less bamboo and leaves. Morphological differences reflect the differences in ecology, habitat, and behavior. Bwindi gorillas measured have smaller bodies, relatively longer limbs, hands, and feet, shorter trunks, thumbs, big toes, and tooth row lengths, and narrower trunks and orbital breadths than Virunga gorillas. These differences indicate Bwindi gorillas do not belong to G.g. beringei and should not be referred to as “mountain gorillas.” How unique the distinguishing features of Bwindi gorillas are, and whether or not they should be assigned to a new taxon, depends on the expression of these features in eastern lowland gorillas (G.g. graueri). © 1996 Wiley-Liss, Inc.     

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.     

Insectivory by Gorilla gorilla gorilla in Southeast Cameroon

Our study extends quantitative analyses of insect-eating by gorillas (Gorilla gorilla gorilla) to Cameroon. During a 2-mo period (May–June 2001), we recorded and analyzed feeding traces on plants and insects and in gorilla feces. We found 180 feeding traces, 17% of which involved insectivory. Seventy-eight percent of the fecal samples (n = 36) contained insects. Ants were found in 61% of the samples, termites in 39%, while 56% of the samples contained remains of other insects. We added 14 new species to the known insect diet of western lowland gorillas. Overall, social insects are predominant. The choice of prey by the Ntonga gorillas gives new clues for the existence of cross-cultural differences among gorilla populations. A comparison of the overall frequency of insectivory with those at other sites in Central Africa indicate a possible effect of forest disturbance on the insectivorous behavior of gorillas. The study suggests the existence of temporal variation in ant- and termite-eating by gorillas.     

Medical Survey of the Local Human Population to Determine Possible Health Risks to the Mountain Gorillas of Bwindi Impenetrable Forest National Park,Uganda

There has been increasing contact between mountain gorillas (Gorilla gorilla beringei) and the human population surrounding Bwindi Impenetrable Forest National Park (BIFNP) in Uganda. Due to the close taxonomic relationship between humans and gorillas there is potential for disease transmission between them. Preventing the introduction or spread of transmissible diseases to the gorillas is essential to protect them. We interviewed 301 villagers living in close proximity to BIFNP with a medical questionnaire in July, 2000. We collected information on demographics, vaccination and health history, and human/gorilla interaction. Our objectives were to estimate the prevalence of several diseases in the human population and to evaluate the risk of anthropozoonotic transmission from humans to gorillas. We found a high prevalence of disease symptoms such as coughing (72.1%) and fever (56.1%) compatible with acute infectious diseases; over half of the respondents (59.1%) had a specific disease diagnosis within the 6 mo preceding the study. We compared villagers who had visual contact with gorillas in the 6 mo preceding the study (53.5%) to villagers who had no visual contact (46.5%). Men were 2.3 times more likely than women to have visual contact with gorillas. In general, the frequency of disease history and symptoms was similar for people with and without contact. The high prevalence of acute infectious diseases in the population surrounding BIFNP and the high rate of contact with gorillas creates the potential for anthropozoonotic disease transmission.