Ranging patterns of Japanese macaques in the coniferous forest of Yakushima: Home range shift and travel rate.

Ranging is one of the most important behavioral adaptations for coping with seasonally fluctuating food and thermal conditions. We studied the ranging patterns, in particular home range shift and travel rate of Japanese macaques (Macaca fuscata) in the coniferous forest of Yakushima by tracking a group for 17 months. We also supplemented our data with records collected every August over a 5-year annual census. The macaque group used the eastern part of their home range from May to September and the western part during the rest of the year. The eastern part of the home range was largely primary forest in the national park, and the altitude was higher than in the western part. When they used the western part, the macaques ate more herbs and fruits from small-sized trees, the availability of which was higher in the logged forest. This east-west home range shift occurred repeatedly over multiple years. A neighboring group occupied the western part of the focal group's home range in summer, which the focal group did not use in that season. Both temperature and diet affected seasonal changes in the monthly average travel rate. Animals need more energy for thermoregulation when the temperature is low, so the macaques decreased their ranging efforts to save energy at times of low temperature. They increased their ranging distance to eat fungi, since their encounters with this food would increase with the total distance walked. They also increased their travel rate when eating flowers, which had lower food patch (tree) density than other foods such as fruits. The data supported the hypothesis that the macaques capitalize on habitat heterogeneity in a seasonally fluctuating habitat by shifting their home range and modifying their travel rate.

Food conditions, competitive regime, and female social relationships in Japanese macaques: within-population variation on Yakushima.

Feeding conditions, competitive regime, and female social relationships of Japanese macaques (Macaca fuscata) on Yakushima were compared between the two habitats at two different altitudes (coniferous forest, 1,000-1,200 m and coastal forest, 0-200 m). Fruit availability was higher in the coastal forest. There was no consistent difference in the frequency of agonistic interactions within a group during feeding between the two habitats. The coastal forest evoked stronger inter-group contest competition compared to the coniferous forest as evidenced by a higher inter-group encounter rate and a higher proportion of aggressive encounters to non-aggressive ones. Birth rate was higher in larger groups compared to smaller ones in the coastal forest, but did not differ in the coniferous forest. In spite of these differences in competitive regime, no variation in female social relationships was observed, such as direction and concentration on particular individuals in grooming, linearity in dominance rank, counter-attack, and support of juvenile kin during agonistic interactions. The present results indicate that the female social relationships of Japanese macaques are robust and do not change according to changes in the current environment.

Behavioral thermoregulation of wild Japanese macaques: comparisons between two subpopulations.

We studied the behavioral thermoregulation of Japanese macaques in two troops that live in the coniferous (1,000-1,200 m in elevation) and coastal forests (0-200 m in elevation) of Yakushima. Frequency of sunbathing, huddling, and microhabitat selection during inactivity was compared. The difference in mean annual air temperature between the forests was more than 7 degrees C. In both forests, when the weather was clear, macaques spent more time being inactive in the sunshine in winter than in autumn. In winter, they huddled more often when it was clear than when cloudy. Microhabitat selection to stay in the sunshine during winter differed between the two forests. In winter, macaques spent more time inactive in open habitats in the coniferous forest and in the trees in the coastal forest than in autumn, respectively. This difference is related to the lower crown height in the coastal forest and the large open habitats (logged area) available only in the coniferous forest. In winter, skin temperature measured by temperature-sensitive transmitters was 1.32-1.71 degrees C higher when sunbathing, and 0.83-4.75 degrees C higher when huddling than staying in the shade without huddling. In winter, the proportion with which they stayed in the sunshine or huddled in winter did not differ between the two forests, in spite of the difference in air temperature. This suggests that Japanese macaques respond to seasonal changes in air temperature, not the absolute temperature, and that they acclimatize themselves to thermal conditions that require behavioral thermoregulation only during the season when thermoregulation is most costly.

Postglacial population expansion of Japanese macaques (Macaca fuscata) inferred from mitochondrial DNA phylogeography.

We investigated the diversity and phylogeography of mitochondrial DNA (mtDNA) in Japanese macaques (Macaca fuscata), an endemic species in Japan that has the northernmost distribution of any non-human primate species. DNA samples from 135 localities representing the entire range of this species were compared. A total of 53 unique haplotypes were observed for the 412-bp partial mtDNA control region sequence, with length variation distinguishing the two subspecies. Clustering analyses suggested two putative major haplogroups, of which one was geographically distributed in eastern Japan and the other in western Japan. The populations in the east showed lower mtDNA diversity than those in the west. Phylogeographical relationships of haplotypes depicted with minimum spanning network suggested differences in population structure. Population expansion was significant for the eastern but not the western population, suggesting establishment of the ancestral population was relatively long ago in the west and recent in the east. Based on fossil evidence and past climate and vegetation changes, we inferred that the postulated population expansion may have taken place after the last glacial period (after 15,000 years ago). Mitochondrial DNA showed contrasting results in both variability and phylogenetic status of local populations to those of previous studies using protein variations, particularly for populations in the periphery of the range, with special inference on habitat change during the glacial period in response to cold adaptation.

Low genetic diversity and biased distribution of mitochondrial DNA haplotypes in the Japanese macaque (Macaca fuscata yakui) on Yakushima Island.

The Japanese macaques (Macaca fuscata yakui) on Yakushima Island are an endemic subspecies and are closely related to the population of Kyushu, one of the main islands of Japan. Using feces collected throughout Yakushima Island, we examined mitochondrial DNA (mtDNA) to investigate the phylogeography of Japanese macaques. Six haplotypes were observed for a 203-bp fragment of the mtDNA control region. The nucleotide diversity (pi) was low (0.0021). The genetic divergence within the Yakushima population was lower (0.009) than that among four haplotypes of the Kyushu population (0.015), calculated using Kimura's two-parameter method. The mismatch distribution analysis of the six haplotypes of the Yakushima population suggested that the Yakushima population had experienced a sudden expansion in population size, which could be related to the bottleneck effect. The geographic distribution of the mtDNA haplotypes was not uniform. One haplotype was distributed widely, whereas the other five haplotypes were distributed only in the lowlands. The low genetic diversity and biased distribution are discussed in relation to an environmental crash caused by ancient volcanic activity near this island, which is postulated to have happened about 7,300 years ago, and the delayed recovery of highland vegetation.

Comparisons of food availability and group density of Japanese macaques in primary, naturally regenerated, and plantation forests.

We compared food availability and group density of Japanese macaques in Yakushima, southern Japan, among primary forest and two habitats that had been disturbed by logging and had different regeneration histories. The study was conducted in an undisturbed national park, forest that was logged 7-18 years ago and later naturally regenerated, and forest that was logged 19-27 years ago and later planted with Japanese cedar (Cryptomeria japonica) trees. The plantation forest was primarily composed of large Cryptomeria japonica trees at low stand density, while the naturally regenerated forest was composed of many small trees. The total basal area and number of trees in the primary forest were comparable to those in the plantation forest. Annual fruit production was greatest in the naturally regenerated forest, intermediate in the primary forest, and negligible in the plantation forest. Herb availability was high in the naturally regenerated forest, but low in the primary and plantation forests. The group density of Japanese macaques was high in the naturally regenerated forest, intermediate in the primary forest, and low in the plantation forest. Since group size in the naturally regenerated forest was small, individual density was almost the same as in the primary forest. These results suggest that the effects of regeneration on macaques vary between the two habitats. The plantation forest consisted mostly of Cryptomeria japonica, which supplies only flowers as food in a limited season, and had a lower density of macaques. On the other hand, in the naturally regenerated forest, fruit production and herb availability were high (probably because of the enhanced light conditions after logging), and the density of macaques was as high as in the primary forest.