Juichi Yamagiwa

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 groups 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.

Reproduction of Wild Japanese Macaque Females of Yakushima and Kinkazan Islands: A Preliminary Report

Wild Japanese macaque females of the Yakushima and Kinkazan populations exhibited similar reproductive features. (1) Births/female/year (BR: 0.27–0.35) was lower than those of provisioned troops, but (2) infant mortality (IM: 0.23–0.25) was higher than those of provisioned troops. (3) The interbirth interval (IBI) following the death of infants was 1.5–1.6 years, shorter than that following surviving infants (2.2–2.4 yrs). (4) Birth sex ratio (BSR) did not differ from 1∶1. There was no consistent correlation between (5) female age and IM, (6) maternal rank and offspring BSR, or (7) maternal rank and reproductive success. On the other hand, (8) BR of Yakushima females was significantly lower than that of Kinkazan females. In particular, (9) Yakushima females stopped reproduction earlier than Kinkazan females, although (10) the first birth of Yakushima females was about one year earlier than Kinkazan females. (11) BR exhibited a humped curve against female age in Yakushima, but it was uncertain whether old-aged females of Kinkazan exhibited a post-reproductive life span (PRLS). (12) The survivorship for female juveniles was lower than that for male juveniles in Yakushima, whereas the survivorship for male juveniles was lower than that for female juveniles in Kinkazan. These data may indicate that Yakushima females more severely compete for resources than Kinkazan females, because of high population density, whereas the population density of Kinkazan might be limited by climate (e.g. heavy snow) rather than density dependent ecological effects.

Seasonal change in the composition of the diet of eastern lowland gorillas

Details are presented of the composition of the diet of eastern lowland gorillas, derived mainly from a study of their fresh trails and fecal analysis, during the course of an entire year in the tropical lowland forests of the Itebero region, Zaire. Gorillas ate 194 plant foods from 121 species and 45 families. They consumed 48 species of fruits; and 89% of fecal samples contained fruit seeds, but fruits were a relatively small part (25%) of the total number of food items. The composition of their diet changed seasonally. When consumption of fruit decreased in the long rainy and the long dry seasons, the gorillas ate, in addition to Zingiberaceae and Marantaceae, many kinds of leaf and bark, which may be an important buffer against the shortage of fruits. Gorillas also fed regularly on ants (Ponerinae), and the frequency of consumption showed small seasonal variations. From a comparison of diet composition, eastern lowland gorillas appeared to be intermediate between the other two subspecies. The choice of food showed differences in preference of fruits and insects between subspecies and may reflect high similarities within subspecies in lowland and montane forests.

Socioecological factors influencing population structure of gorillas and chimpanzees.

Differences in distribution and density between gorillas and chimpanzees are reconsidered with special reference to population structure. Both ecological and social factors influencing population structure are compared between species and between habitats within species. Gorillas and chimpanzees respond differently to a decline in food quality, such as fruit scarcity: gorillas change diet and decrease range, while chimpanzees do not change diet but may expand range. These responses result in different effects on their grouping patterns. For gorillas the dispersed distribution and reduction of range size decreases the rate of inter-unit encounters and female transfer. The concentration of social units increases the rate of aggressive contact between units and stimulates female transfer. Social units of gorillas may crowd or disperse in order to attain the optimal density. This tendency may result in similar densities of gorillas across habitats. By contrast, the distribution patterns or range size may not affect inter-unit relationships of chimpanzees. Within a single unit-group, various reproductive strategies are adopted by both sexes. Independent travel of females and flexible grouping patterns enable them to survive at very low density in extraordinary large ranges. Density and inter-unit relationships are good criteria for a healthy population of gorillas, while the size of unit-group and inter-individual relationships are good criteria for chimpanzees. Conservation planners should consider these differences for sympatric and allopatric survival in these species.

Factors Affecting Nesting Site Choice in Chimpanzees at Tshibati, Kahuzi-Biega National Park: Influence of Sympatric Gorillas

We recorded 310 fresh chimpanzee night nests at 72 nest sites to determine their choice of tree and site for nesting vis-à-vis the effects of sympatric gorillas. Chimpanzees did not use trees for nesting according to their abundance, but instead tended to nest in fruit trees that they used as food sources. Nesting patterns of chimpanzees may vary with nesting group size, the type of vegetation, and fruit species eaten or not eaten by gorillas. When chimpanzees lodged as a small group in the secondary forest, they nested more frequently in trees bearing ripe fruits eaten only by themselves than in those with fruit eaten also by gorillas. When they lodged as a large group in the primary forest, they nested more frequently in trees bearing ripe fruits eaten by both apes. Nest group size is positively correlated with the availability of preferred ripe fruits in secondary forest. These findings not only reflect the larger foraging groups at the larger fruiting trees but also suggest that chimpanzees may have tended to occupy fruiting trees effectively by nesting in them and by forming large nest groups when the fruits attracted gorillas. Competition over fruits between gorillas and chimpanzees, due to their low productivity in the montane forest of Kahuzi, may have promoted the chimpanzee tactics.

A census of the eastern lowland gorillas Gorilla gorilla graueri in Kahuzi-Biega National Park with reference to mountain gorillas G. g. beringei in the Virunga Region, Zaire

Abstract A population census of eastern lowland gorillas Gorilla gorilla graueri was conducted in the original part of Kahuzi-Biega National Park, Zaire. At least 258 gorillas (25 groups and nine solitary males) were estimated by bed counts to have survived within the Park in the 1990 bamboo season. In comparison with results of the previous censuses, conducted between 1978 and 1979, the population showed a slight increase. The percentage of immature individuals in the population indicates that gorillas still maintain a healthy population, but mean group size has decreased markedly. Although gorillas expanded their range in the 11 years between the surveys, they have still tended to concentrate in the central well-protected area of the Park, possibly stimulating frequent transfer of females between groups or from groups to solitary males, resulting in the decrease in group size. Recent social changes recorded from observations of habituated groups tend to support this interpretation. Human disturbance in the bridge zone between highland and lowland forests prevents gorillas from making contact with their neighbours and reduces the possibility of outbreeding within the Park. Reference is made to the more detailed censuses of the mountain gorilla G. g. beringei . More international attention will be needed to increase conservation activity to protect gorillas from the hazards of human disturbances in this area.

How fruit abundance affects the chimpanzee party size: a comparison between four study sites

We examined the relationship between fruit abundance and chimpanzee (Pan troglodytes) party size by comparing data from four study sites: the Kalinzu Forest Reserve, Uganda, the Djinji Camp and Guga Camp in the Ndoki Forest, Congo, and Kahuzi-Biega National Park, Democratic Republic of Congo. Although the difference in the fruit abundance between the sites was responsible for the difference in the party size between the sites, the seasonal changes in fruit abundance did not explain the changes in the party size in each study site. Across the four study sites, there were significant correlations of the mean and minimum of monthly party size with the mean of monthly fruiting-tree density, and a significant correlation of the maximum of monthly party sizes with the minimum of monthly fruiting-tree density. We proposed a hypothesis that (1) the monthly fruit abundance affects the monthly party size in the sites where the fruit availability is as low as to limit the party size during a major part of a year, while (2) the party size does not increase with the increase in the monthly fruit abundance, but is affected by other social factors, in the sites where the minimum of monthly fruit abundance is high enough for chimpanzees to form parties of an adequate size.

Fallback foods and dietary partitioning among Pan and gorilla

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

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 groups 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.

Why dominants do not consistently attain high mating and reproductive success: A review of longitudinal Japanese macaque studies.

There is wide interest in the effects of reproductive biology, mating partner preference, and rank on mating success (MS) and reproductive success (RS) in primates. In particular, theory stresses importance on the mechanisms for attaining RS. Most theory hedges on competitive ability and priority of access to resources, whether they be food or estrous females. However, the majority of data used in favor of such hypotheses come from relatively short-term studies. We review these hypotheses based on long-term data from provisioned and unprovisioned populations of Japanese macaques. Neither MS nor RS were consistently attained by high-ranking males and females. For males, female choice and mating partner preference is seen to over-ride most male-male competitive behaviors likely to affect MS and RS through priority of access to estrous females. Long-term mating patterns driven largely by female partner preferences, results in decreasing MS and RS for older higher-ranking males. The long-term trend for females to prefer less familiar or novel partners results in higher MS and RS for younger, middle-ranking males. The effects of this vary according to troop size and the duration of male tenure. For females, no consistent trend was recognized for rank related RS in either provisioned or unprovisioned troops. Non-reproductive mating may provide differential benefit to high-ranking females for access to limited food resources in some habitats but overall the relationship was inconclusive. Distribution and defendability of food resource, rather than provisioning per se may be more important.