No abstract

Seasonal variations in the activity budget of Japanese macaques in the coniferous forest of Yakushima were studied over the course of 1 year. On an annual basis, they spent 38% of the daytime feeding, 16% traveling, 14% in social interactions, and 32% engaged in resting. The effects of temperature and food-related factors (i.e., food distribution, feeding speed, and food abundance) on the seasonal variations of activity budget were examined by stepwise multiple regression analysis. When the temperature was low, the macaques decreased traveling and feeding time, in accordance with the prediction that endothermal animals save energy under severe thermoregulatory cost. When the feeding speed of available foods was slow, they spent more time feeding. When high-quality foods were abundant, they decreased feeding time. These macaques did not respond to fluctuations in food distribution. The present results indicate the importance of temperature, in addition to food-related factors, as a determinant of activity budgets.

We analyzed regional variation in the diets of two primate clades, Asian macaques and colobines, whose distributions include temperate-alpine forests. We addressed feeding strategies that enabled them to adapt to harsh environments characterized by relatively low mean temperatures and strong seasonality in both temperature and food availability. Macaques in tropical-lowland forests feed mainly on fruit and animal matter whereas populations in temperate-alpine forests feed more on foliage and on such items as bark and fungi. In comparison, colobines in tropical-lowland forests feed more on fruit and foliage whereas populations in temperate-alpine forests feed less on flowers and more on lichens. Annual precipitation and mean temperature, both of which reflect primary production, had the most significant effects on the feeding behavior of the macaques, whereas only mean temperature had a significant effect on that of colobines. We found two behavioral strategies used by both clades to cope with severe environmental conditions in temperate-alpine forests--shifting to other food items and adjusting feeding plasticity for fruit and foliage. Macaques responded to latitudinal changes by use of both strategies whereas the colobines adapted by using the latter only. By contrast, changes in altitude resulted in the macaques' using the latter strategy and colobines' using both. The different current distributions of Asian macaques and colobines could be attributed to differences in their feeding strategies originating in their digestive systems.

The orangutan is the world's largest arboreal mammal, and images of the red ape moving through the tropical forest canopy symbolise its typical arboreal behaviour. Records of terrestrial behaviour are scarce and often associated with habitat disturbance. We conducted a large-scale species-level analysis of ground-based camera-trapping data to evaluate the extent to which Bornean orangutans Pongo pygmaeus come down from the trees to travel terrestrially, and whether they are indeed forced to the ground primarily by anthropogenic forest disturbances. Although the degree of forest disturbance and canopy gap size influenced terrestriality, orangutans were recorded on the ground as frequently in heavily degraded habitats as in primary forests. Furthermore, all age-sex classes were recorded on the ground (flanged males more often). This suggests that terrestrial locomotion is part of the Bornean orangutan's natural behavioural repertoire to a much greater extent than previously thought, and is only modified by habitat disturbance. The capacity of orangutans to come down from the trees may increase their ability to cope with at least smaller-scale forest fragmentation, and to cross moderately open spaces in mosaic landscapes, although the extent of this versatility remains to be investigated.

We examined the basal area of two life forms (conifers vs. broadleaf trees) along elevational gradients on Yakushima Island, Japan and on two series of geological substrate on Mount Kinabalu, Borneo. On Yakushima, total stand basal area abruptly increased from 700 to 1,050 m in accordance with the high dominance of conifers, indicating the presence of additive basal area of conifers in conifer-broadleaf mixed forests at higher elevations (1,050-1,300 m). Along two substrate series on Kinabalu, some forests at higher elevations (1,860-3,080 m) showed relatively high dominance of conifers, but conifer basal area did not appear to be additive. Conifers were emergents above the canopy of broadleaf trees in mixed forests on Yakushima, but two life forms usually coexisted in the single-story canopy in mixed forests on Kinabalu. Litterfall rate as a surrogate of productivity decreased with decreasing temperature along elevation on both the sites, but the rate of decrease was slower on Yakushima, where mixed forests at higher elevations showed relatively high rates. Thus, we suggest that additive basal area of conifers was linked to their emergent status, and that it enhanced productivity by complementary use of light by two life forms that occupy different stories. On Yakushima, typhoons are a major disturbance, but do not severely limit the height growth of conifers, allowing the development of twostory mixed forests. On Kinabalu, a major disturbance is El Nin˜o-driven drought, and hydraulic limitation to tree height may explain the non-additive and non-emergent nature of conifers.

Altitudinal variations in relative group densities of the Japanese macaques on Yakushima were studied. This is an ideal place for studying resource limitations because it avoids various complicating factors that are difficult to quantify but might affect animal densities, such as predation, interspecific competition, and past catastrophes. The relative group density was high in the coastal forest (0-400 m), while it did not differ among the higher zones (400-800, 800-1200 and 1200-1886 m). To examine this variation, three habitat variables were analyzed: total basal area of food trees per unit area, seasonal variations in fruit abundance, and total annual fleshy fruit production. All of these variables indicate that fruit and seeds are most available in the coastal forest. Thus, altitudinal variations in the density of Japanese macaques on Yakushima are determined by the total annual food abundance.

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.

We conducted observations of two neighboring groups of food-enhanced long-tailed macaques (Macaca fascicularis) over a period of one year. We examined their diet, behavior, habitat use, and ranging and how within-population variability reflected differential utilization of anthropogenic food resources. The group that consumed more anthropogenic food spent less time feeding on wild fruits and flowers, less time resting, and more time locomoting. They used forest habitats less often, and had a larger total home range and mean monthly home range. Some of these results contrasted with previous studies of food-enhanced primates which reported that food-enhancement resulted in smaller home ranges, shorter daily ranges, less time traveling and feeding, and more time resting. These contrasting patterns may relate to the nature of anthropogenic foods. In most studies of food-enhanced primates, anthropogenic food resources were abundant and concentrated but the macaques in this study used anthropogenic foods mainly from a few refuse sites where they had limited access, and from dispersed and irregular human provisioning. The group consuming more anthropogenic food therefore showed more spatially dispersed feeding activity and home range use, an effect that was likely further enhanced by lower natural food resource availability within their home range. The Singapore macaque population shows small-scale variability in feeding and ranging behavior, contributing to the complexity of their adaptive variability in a human-altered habitat. Our findings could have important implications for mitigating human-macaque conflict as measures applied at a higher spatial or population level may achieve highly inconsistent results, intensifying the challenges for wildlife managers.