Group size and structure in western lowland gorillas (Gorilla gorilla gorilla) at Mbeli Bai, Republic of Congo

This paper describes the size and structure of western lowland gorilla groups visiting Mbeli Bai, a swampy forest clearing in northern Congo. An observation platform at the edge of the clearing was used over a 3-yr period, with gorillas visible for 1,681 hr. Data are presented on 14 groups and seven solitary males. Mean group size (excluding solitary males) was 8.4 +/- SD 4.3, and did not differ significantly from most other gorilla studies. All groups at Mbeli contained only one fully mature male and did not show evidence of fission-fusion or regular subgrouping. All emigrating males that remained in the population became solitary, and "bachelor" groups were not observed. Methodological issues are raised, including the classification of multi-male groups, and the demographic profiles of other populations are discussed in the light of results from Mbeli.     

Mineral content as a basis for food selection by western lowland gorillas in a forest clearing

The forests in northwest Republic of Congo contain a number of herbaceous swamp clearings that provide foraging sites for lowland gorillas (G.g. gorilla). A 10-month study at the Maya Nord clearing (Parc National d'Odzala) showed that feeding activities occupied 72% of the time visiting gorillas spent on the clearing. They fed on four plant species: Enydra fluctuans (Asteraceae), Cyperus sp., Pycreus mundtii, and Rhynchospora corymbosa (Cyperaceae) among the 45 species recorded on the clearing. These clearing food species have higher mineral contents (especially Na and Ca) than the dominant Marantaceae species (Haumania liebrechtsiana) that constituted a staple food plant for gorillas in this forest. They also have higher potassium contents and contain less lignin than non-eaten clearing items/species. Finally, the most actively searched for clearing food (Enydra fluctuans) was characterized by the highest amount of Na and Ca. These results suggest that the mineral content (especially in Na, Ca, and/or K) could determine the feeding selectivity of gorillas at the clearing. They also tend to confirm that the amount of fiber plays a deterrent role in food selectivity, as has been found by many authors. The high density of gorillas in that region could result from the combination of the large areas of Marantaceae forests that provide abundant though monotonous food, and the number of clearings that provide sufficient mineral supplies. Clearings should thus be considered as key habitats for the conservation of gorillas.     

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.     

Population and group structure of western lowland gorillas (Gorilla gorilla gorilla) at Lokoué, Republic of Congo

During a 17-month study at the Lokoué clearing in Odzala National Park, Republic of Congo, we identified 377 western lowland gorillas. This population included 31 solitary males, 37 breeding groups, and eight nonbreeding groups. Its age- and sex-class structure was similar to those observed at two other clearings in the same forest block. However, the size of breeding groups varied with site (either clearing or forest sites). At Lokoué, breeding groups (mean size: 8.2 gorillas; range: 3-15) included a single silverback male and, on average, 3.2 adult females. Nonbreeding groups (mean size: 5.5; range: 2-15) were devoid of adult females. Five of the nonbreeding groups were composed predominantly of blackbacks, subadult males, and juveniles, and thus fit the definition of all-male groups previously observed in mountain gorillas. Our study confirms that 1) one-male breeding groups are the norm in western gorillas, and 2) all-male groups occur in this species. Despite frequent changes in members due to migrations of the males, the persistence of these all-male groups indicates that they may play an important role in the life of migrating males. Variations in population structure, and group composition and type among gorilla populations are discussed. However, a further understanding of the evolution of group-living in gorillas requires detailed ecological studies conducted in parallel with studies of the population structure and dynamics of these groups.     

Survey of Grauer's Gorillas (Gorilla gorilla graueri) and Eastern Chimpanzees (Pan troglodytes schweinfurthi) in the Kahuzi-Biega National Park Lowland Sector and Adjacent Forest in Eastern Democratic Republic of Congo

We describe the distribution and estimate densities of Grauer's gorillas (Gorilla gorilla graueri) and eastern chimpanzees (Pan troglodytes schweinfurthi) in a 12,770-km ² area of lowland forest between the Lowa, Luka, Lugulu, and Oku rivers in eastern Democratic Republic of Congo, the site of the largest continuous population of Grauer's gorillas. The survey included a total of 480 km of transects completed within seven sampling zones in the Kahuzi-Biega National Park lowland sector and adjacent Kasese region and approximately 1100 km of footpath and forest reconnaissance. We estimate total populations of 7670 (4180–10,830) weaned gorillas within the Kahuzi-Biega lowland sector and 3350 (1420–5950) individuals in the Kasese survey areas. Within the same area, we estimate a population of 2600 (1620–4500) chimpanzees. Ape nest site densities are significantly higher within the Kahuzi-Biega lowland sector than in the more remote Kasese survey area in spite of a significantly higher encounter rate of human sign within the lowland sector of the park. Comparison of our data with information obtained by Emlen and Schaller during the first rangewide survey of Grauer's gorillas in 1959 suggests that gorilla populations have remained stable in protected areas but declined in adjacent forest. These findings underscore the key role played by national parks in protecting biological resources in spite of the recent political and economic turmoil in the region. We also show that forest reconnaissance is a reliable and cost-effective method to assess gorilla densities in remote forested areas.     

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.     

Day-journey length and daily diet of solitary male gorillas in lowland and highland habitats

We compared day-journey length and daily diets of solitary male gorillas in lowland versus highland habitats. Solitary males in tropical forests of Zaire tend to travel longer distances, to visit more types of vegetation, and to consume more kinds of food than a solitary male mountain gorilla in the Virunga Volcanoes did. The number of feeding sites per day is larger and the mean distance between feeding sites is far longer for the former than the latter. These observations may reflect differences in food breadth and availability between highland and lowland habitats. The herbaceous plants that are eaten by mountain gorillas are densely and evenly distributed in the higher montane forest of the Virungas, where gorillas need not cover long distances to search for food. In contrast, herbaceous plants are scarce in primary and ancient secondary forests of lowland habitats, where gorillas travel long distances and eat various fruits and insects. The patchy and unpredictable distribution of foods may extend the distances over which gorillas search for food in the lowland habitat. However, solitary males showed a prominent reduction in day-journey length and changed their choices of food during the nonfruiting season (the long rainy and dry seasons) in the lowland habitats. This strategy may have developed during the Pleistocene and may have enabled them to enlarge their ranges to the higher montane forests, where fruits are sparse throughout the year.     

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.     

Recovery potential of a western lowland gorilla population following a major Ebola outbreak: results from a ten year study

Investigating the recovery capacity of wildlife populations following demographic crashes is of great interest to ecologists and conservationists. Opportunities to study these aspects are rare due to the difficulty of monitoring populations both before and after a demographic crash. Ebola outbreaks in central Africa have killed up to 95% of the individuals in affected western lowland gorilla (Gorilla gorilla gorilla) populations. Assessing whether and how fast affected populations recover is essential for the conservation of this critically endangered taxon. The gorilla population visiting Lokoué forest clearing, Odzala-Kokoua National Park, Republic of the Congo, has been monitored before, two years after and six years after Ebola affected it in 2004. This allowed us to describe Ebola's short-term and long-term impacts on the structure of the population. The size of the population, which included around 380 gorillas before the Ebola outbreak, dropped to less than 40 individuals after the outbreak. It then remained stable for six years after the outbreak. However, the demographic structure of this small population has significantly changed. Although several solitary males have disappeared, the immigration of adult females, the formation of new breeding groups, and several birth events suggest that the population is showing potential to recover. During the outbreak, surviving adult and subadult females joined old solitary silverbacks. Those females were subsequently observed joining young silverbacks, forming new breeding groups where they later gave birth. Interestingly, some females were observed joining silverbacks that were unlikely to have sired their infant, but no infanticide was observed. The consequences of the Ebola outbreak on the population structure were different two years and six years after the outbreak. Therefore, our results could be used as demographic indicators to detect and date outbreaks that have happened in other, non-monitored gorilla populations.     

Recent divergences and size decreases of eastern gorilla populations

Compared with other African apes, eastern gorillas (Gorilla beringei) have been little studied genetically. We used analysis of autosomal DNA genotypes obtained from non-invasively collected faecal samples to estimate the evolutionary histories of the two extant mountain gorilla populations and the closely related eastern lowland gorillas. Our results suggest that eastern lowland gorillas and mountain gorillas split beginning some 10 000 years ago, followed 5000 years ago by the split of the two mountain gorilla populations of Bwindi Impenetrable National Park and the Virungas Massif. All three populations have decreased in effective population size, with particularly substantial 10-fold decreases for the mountain gorillas. These dynamics probably reflect responses to habitat changes resulting from climate fluctuations over the past 20 000 years as well as increasing human influence in this densely populated region in the last several thousand years.     

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.     

Use of buckets as tools by Western lowland gorillas

While all great apes have been documented to use tools, gorillas are arguably the least proficient tool users. In 2009, a Western lowland gorilla (Gorilla gorilla gorilla) at the Buffalo Zoo was observed using a bucket, which had been provided as part of normal enrichment, as a tool to collect water. We conducted a brief, ad libitum investigation to confirm the validity of the initial observation. We then carried out a systematic investigation of the behavior in 2010. We collected 72 hr of videotaped data and tested the null hypothesis that the gorillas did not differ in their prevalence of engaging in bucket-use behaviors. We documented that all four adult gorillas in the group used buckets as drinking tools; however, there was significant individual variation in frequency and type of use of buckets. Four of the eight behaviors showed significant variation among individuals. The silverback male and the youngest adult female contacted and held the bucket significantly more than the remaining two adult females. The young female carried and drank from the bucket significantly more than any other individual. Furthermore, she was observed to fill the bucket with water four of the six times during which this behavior was observed. These data provide evidence of the ability of gorillas to utilize tools, given the appropriate environmental conditions. We continue to explore the abilities of gorillas to recognize the functionality of buckets as tools.     

Genetic diversity of North American captive‐born gorillas (Gorilla gorilla gorilla)

Western lowland gorillas (Gorilla gorilla gorilla) are designated as critically endangered and wild populations are dramatically declining as a result of habitat destruction, fragmentation, diseases (e.g., Ebola) and the illegal bushmeat trade. As wild populations continue to decline, the genetic management of the North American captive western lowland gorilla population will be an important component of the long‐term conservation of the species. We genotyped 26 individuals from the North American captive gorilla collection at 11 autosomal microsatellite loci in order to compare levels of genetic diversity to wild populations, investigate genetic signatures of a population bottleneck and identify the genetic structure of the captive‐born population. Captive gorillas had significantly higher levels of allelic diversity (t₇ = 4.49, P = 0.002) and heterozygosity (t₇ = 4.15, P = 0.004) than comparative wild populations, yet the population has lost significant allelic diversity while in captivity when compared to founders (t₇ = 2.44, P = 0.04). Analyses suggested no genetic evidence for a population bottleneck of the captive population. Genetic structure results supported the management of North American captive gorillas as a single population. Our results highlight the utility of genetic management approaches for endangered nonhuman primate species.     

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.     

Living in nonbreeding groups: an alternative strategy for maturing gorillas

The one-male reproductive strategy implies that maturing males are temporarily excluded from reproduction. In gorillas, these excluded males live either solitarily or in nonbreeding groups (NBGs) that are devoid of adult females. The dynamics of NBGs are not well known. In this study, which was conducted on a gorilla population (Gorilla gorilla gorilla) of 377 individuals that visited the Lokoué clearing in the Republic of Congo, we detail how the NBGs formed, and analyze their dynamics according to age-sex classes, the relatedness of members, and the origin and destination of transferring individuals. We discuss the potential benefits gained by individuals living in these groups. The NBGs included mainly immature males, most of which appeared to have migrated voluntarily from their natal groups. Some individuals (including juvenile females) came from disbanded breeding groups (BGs). Migrants preferentially joined NBGs that included a silverback male. Their dispersal patterns were not determined by their degree of relatedness, but they tended to associate with related silverbacks. In this way, the migrants could enhance their protection against predators and gain experience with different environmental conditions. By tolerating and protecting offspring, aging silverbacks could enhance their inclusive fitness. Finally, young and healthy silverbacks could increase their likelihood of forming a future BG when unrelated females joined them.     

Modeling the Impact of Ebola and Bushmeat Hunting on Western Lowland Gorillas

The 2003 outbreak of Ebola in the Republic of Congo killed 114 people and up to 800 western lowland gorillas. This outbreak and all outbreaks between 2001–2003 began with human handling of infected animal carcasses. Ebola has since spread, putting the entire gorilla population at risk. An epidemiological model is presented to describe the combined effects of Ebola and hunting on persistence of gorillas. The number of infected gorillas also provides a means of assessing the risk of transmission to humans. Under current harvest practices and the estimated annual outbreak rate, the gorilla population is predicted to undergo a 97% decline within 100 years. Controlling bushmeat hunting may not be enough to prevent extinction if frequent outbreaks occur.     

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.     

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.     

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.     

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.