Postconflict behavior in captive western lowland gorillas (Gorilla gorilla gorilla)

Postconflict (PC) behaviors, including reconciliation and consolation, have been observed in many primate and several nonprimate species. Using the PC-matched control (MC) method, PC behavior was examined in two groups (n=13) of captive western lowland gorillas, a species for which no conflict resolution data have been published. Analyses of 223 conflicts showed significantly more affiliation between former opponents after a conflict when compared to control periods, indicating reconciliation. Results also showed significantly more affiliation between the victim and a third-party after a conflict, indicating consolation. Both solicited and unsolicited consolation were observed. The majority of the affiliative interactions observed for both reconciliation and consolation were social proximity, which suggests that unlike most nonhuman primates, proximity, rather than physical contact, may be the main mechanism for resolving conflicts in western lowland gorillas. PC behavior was not uniform throughout the groups, but rather varied according to dyad type.     

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.     

Peaceful post-conflict interactions between aggressors and bystanders in captive lowland gorillas (Gorilla gorilla gorilla)

We observed a colony of lowland gorillas in order to assess the occurrence of Peaceful Post-Conflict Interactions between Aggressors and Bystanders (PPIAB). We compared the dynamics of PPIAB with those of Peaceful Post-Conflict Interactions between Victims and Bystanders (PPIVB), which are directed toward victims. We confirmed the occurrence of PPIAB (mean Triadic Contact Tendency: 41.7%+/-16.2% SEM) at frequencies comparable to PPIVB. Immature gorillas were responsible for most PPIAB. PPIAB occurred when the bystander was unrelated to either the aggressor or the victim, and when it was a relative of the aggressor. However, affinitive contact rates were higher in the former case. Unlike PPIVB, PPIAB significantly reduced the likelihood of further aggressions within the group, suggesting that they work at an immediate level. Our data also suggest that bystanders decide when a peaceful contact may be too risky; in fact, neither PPIAB nor PPIVB occurred after high-intensity aggressions.     

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.     

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.     

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.     

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.     

Testicular histological examination of spermatogenetic activity in captive gorillas (Gorilla gorilla)

To clarify the reproductive state of male gorillas, we performed histological examinations on the testicles of 10 male gorillas (Gorilla gorilla). The testicular samples were obtained by autopsy, and ordinal histological preparations were made for light microscopy. The poor spermatogenesis of this species was characterized by the following findings: First, spermatogenesis was evident in only four samples. Meiosis progressed in two samples, but they lacked spermatogenesis. In the remaining four specimens, seminiferous tubules hyalinized without any sign of spermatogenesis. Second, seminiferous epithelia were thin even in the males in which spermatogenesis was observed. Third, degenerated seminiferous tubules were found in all specimens. Fourth, abnormally large syncytial cells were found in the tubules. Six stages in the epithelial cycle of the seminiferous tubules were identified. Testosterone staining made it clear that there were many Leydig cells with spherical or fusiform nuclei in an abundance of interstitial tissue. The relevance of the testicular architecture of gorillas to the mating system is discussed.     

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.     

Intra- and inter-group interactions of an all-male group of virunga mountain gorillas (Gorilla gorilla beringei)

Six unrelated male gorillas formed an all-male group within the Virunga mountain gorilla population. Frequent homosexual interactions characterized the high cohesiveness of this group. Such homosexual behavior reduced the inter-individual distances and increased the social tension between the two silverbacks in the group. The silverbacks retained “ownership” of the homosexual partners, but competed and fought with each other violently when the partners avoided or ignored their courtship. Neither submissive nor reassurance behavior was noted between the silverbacks. Thus, their relationships may not be explained in terms of dominance and subordinancy. However, the loser-support and mediating behavior observed in the group prevented them from engaging in severe fights. Aggression was always directed from the elder and dominant males to the younger and subordinate males, while supporting interactions occurred in the opposite direction. The blackbacks frequently supported the subadult aggressees by attacking the silverbacks, and the younger males displayed mediating behavior in violent fights between the silverbacks. The group's ranging was influenced by encounters with neighboring social units. The members avoided contact with other units and shifted their range after several encounters. On the other hand, when and after a subadult male had immigrated into their group, they frequently encountered other units and did not move away from the encounter site. An all-male group may not be a favorable unit for females to transfer to, but may be profitable for maturing males to associate with. Its formation is probably related to recent social change in the Virunga gorilla population.     

Spatial memory and foraging competition in captive western lowland gorillas (Gorilla gorilla gorilla)

Spatial memory and foraging competition were investigated in three mother/offspring pairs of western lowland gorillas,Gorilla gorilla gorilla, using a naturalistic foraging task at the Toronto Zoo. Sixteen permanent food sites were placed throughout the animals’ enclosures. All of the sites were baited and a pair of animals was free to visit the sites and collect the food. Five of the subjects collected the food with accuracy better than chance. Most of the subjects visited the sites using a pattern, and for half the subjects this was one of adjacency. The high accuracy of five of the subjects and the lack of a consistent adjacency pattern suggest that the animals did in fact use spatial memory. Furthermore, the gorillas tended to avoid visiting food sites that had been previously depleted by their partner. They also appeared to split their search of the enclosures, each visiting only a proportion of the food sites. This indicated that the animals were competing and altering their foraging behaviour based on the behaviour of their partner. Therefore, accuracy was recalculated to take this into account. When the sites depleted by either animal in a pair during a given trial were worked into the accuracy calculations for individual animals, three of the animals still maintained accuracy above chance. This suggests that the animals were not only able to remember which sites they had depleted, but those sites depleted by their foraging partner as well.     

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.     

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.     

Within-group social relationships among females and adult males in wild western lowland gorillas (Gorilla gorilla gorilla)

Within-group female relationships, and relationships between females and adult males have not been described for wild western lowland gorillas. New data are presented here from Mbeli Bai in the Nouabale-Ndoki National Park, Republic of Congo. Nine groups were observed over a continuous 2-year period, and affiliative and agonistic interactions, as well as time spent by females and adult males in proximity to each other were recorded. Affiliative behavior was extremely rare in the bai, and measures of proximity between individuals were inconclusive because of the small sample sizes. Rates of agonistic interaction among females were highly variable between groups. The majority of agonistic interactions were undecided, and the supplant rate was low. Evidence for an agonistic dominance hierarchy is weak. The data do not support hypotheses that increased frugivory in western lowland gorillas raises levels of within-group contest competition; however, the need for intersite comparisons is stressed. In contrast, rates of agonistic behavior between females and silverback males were higher and more consistent between groups. Agonistic relationships between males and females conformed with patterns seen in mountain gorillas, where male protection from intraspecific aggression is important. The benefit females gain by forming long-term cooperative alliances appears to be small, and is likely outweighed by reproductive advantages associated with male protection.     

Activity rhythm and the ranging of a solitary male mountain gorilla (Gorilla gorilla beringei)

To clarify the advantages of solitary life in gorilla males, a lone silverback mountain gorilla (Gorilla gorilla beringei) was studied for nine months in the natural habitat of the Virunga volcanoes. While the time budget for each activity and daily activity cycle were similar to those of groups, his daily journey distance and ranging patterns differed from those of groups. His movements were little influenced by the distribution and abundance of foods, which strongly influence the movements of groups. He notably increased his day journey distances when he encountered neighboring groups. He persistently followed the groups for days and went out of his usual range area. These encounters shifted his monthly range from his natal group's range to that of other groups. When the silverbacks of the encountered groups noticed his presence, they usually gave hoots and chest-beats and sometimes fought violently with him, while females and immatures did not show positive responses towards him. Lone males could have more chance to contact females and to lure them away from their groups than silverbacks within groups. The lone male stage, accompanied by frequent contacts with different groups, probably provides maturing males with useful knowledge of neighboring groups and areas.