Goro Hanya

Diet of a Japanese macaque troop in the coniferous forest of Yakushima

I studied the diet of a troop of Japanese macaques in the coniferous forest of Yakushima over one year via focal animal sampling. Fiber-rich foods constituted 45% of annual feeding time, and mature leaves constituted 38% of total feeding time. Feeding time on fruits and seeds was only 13% and 4%, respectively. Flowers and fungi contributed a considerable amount of annual feeding time: 15% and 14%, respectively. Their diet changed seasonally. They fed on more fruits and seeds in response to increased availability, and when these foods were not available, they ate mature leaves. When the temperature was low, they ate more herbs, possibly to save energy by not climbing trees and staying in sunny places. The results imply two dietary characteristics of the species that might relate to adaptations in temperate regions, where fruit is available during a limited season and fiber-rich foods are the only candidate of fallback food. They are capable of digesting a large amount of fiber-rich food, but at the same time they preferentially select high-quality fruits or seeds when they are available.

Not only annual food abundance but also fallback food quality determines the Japanese macaque density: evidence from seasonal variations in home range size.

Previous studies on Japanese macaque (Macaca fuscata) densities suggest that both total annual food abundance and the quality of fallback foods in the winter bottleneck period affects density. We reviewed data on the seasonal changes in home range size to explain how both factors affect density. In general, home range was large in summer or autumn and small in spring or winter, indicating that density is determined by the home range size in the seasons before winter. The main foods in these seasons are fruits and seeds. If these foods are not abundant, macaques need to range over a larger area, thus decreasing density. Macaques survive the winter by depending on the fat deposited before winter through eating these high-quality foods. If the food condition in winter is severe and the amount of required fat deposition is large, macaques need a larger home range before winter, and thus density becomes lower.

Environmental determinants of the altitudinal variations in relative group densities of Japanese macaques on Yakushima

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.

Feeding strategies of primates in temperate and alpine forests: comparison of Asian macaques and colobines

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.

Vertical Distribution of Wild Yakushima Macaques (Macaca fuscata yakui) in the Western Area of Yakushima Island, Japan : Preliminary Report

A census of wild Yakushima macaques (Macaca fuscata yakui) was carried out in a 23-km2 area of the western coast of Yakushima Island, Japan. We analyzed the census data to investigate changes in monkey distribution associated with the vertical distribution of vegetation. In the lowland coastal zone of 0–300 m above sea level (a.s.l.), 4.8 troops and 62.4–99.8 monkeys are estimated to have existed per km2. In the mountainside zones of 300–900 m a.s.l., the troop density decreased to 1.3–1.6 troops/km2. Since there was no difference in size between the coastal and mountainside troops, population density should decrease with altitude to about 30–36 monkeys per km2. On the other hand, 2.4 troops and about 36 monkeys were estimated to have inhabited per km2 in the mountain summit zone of 900–1,323 m a.s.l.

Linking feeding ecology and population abundance: a review of food resource limitation on primates

We review studies that consider how food affects primate population abundance. In order to explain spatial variation in primate abundance, various correlates that parameterize quality and quantity of food in the habitat have been examined. We propose two hypotheses concerning how resource availability and its seasonality determine animal abundance. When the quality of fallback foods (foods eaten during the scarcity of preferred foods) is too low to satisfy nutritional requirement, total annual food quantity should determine population size, but this relationship can be modified by the quality or the quantity of fallback foods. This mechanism has been established for Japanese macaques and sportive lemurs that survive lean seasons by fat storage or extremely low metabolism. Second, when fallback food quality is high enough to satisfy nutritional requirement but quantity is limited, quantity of fallback food should be a limiting factor of animal abundance. This is supported by the correlation between fig density, which is a high-quality fallback food, and gibbon and orangutan abundance. For a direct test of these hypotheses, we need more research that determines both the quality of food that animals require to satisfy their nutritional requirement and the quantity of food production. Leaves are often regarded as superabundant, but this assumption needs careful examination.

Diet, activity, habitat use, and ranging of two neighboring groups of food-enhanced long-tailed macaques (Macaca fascicularis).

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. Am. J. Primatol. 75:581‐592, 2013.

Comparisons of dispersal success between the species fruiting prior to and those at the peak of migrant frugivore abundance

I evaluated whether plants gain high dispersal success by synchronizing their fruiting with frugivore abundance. Fruiting phenologies, seasonal fluctuations in the abundance of frugivorous birds, and consumption of fruits by birds and Japanese macaques, Macaca fuscata Blyth, were studied in the montane forest of Yakushima for two years. At the community level, fruiting phenologies and seasonal fluctuations in frugivorous bird abundance were asynchronous because Zosterops japonica Temminck et Schlegel, a resident frugivore, decreased in number during the fruiting season. In addition, Symplocos myrtacea Sieb. et Zucc. fruited in September, before the migration of frugivorous birds (Turdus spp. and brown-eared bulbuls Hypsypetes amaurotis Temminck) in November and December. The phenology of other fruit species (Eurya japonica Thunb. and Cleyera japonica Thunb.p.p.emend. Sieb. et Zucc) were synchronized with migrant frugivorous birds. Fruit species with phenologies that are synchronized with migrant frugivore abundances have higher dispersal success either by birds (C. japonica) or macaques (E. japonica). Macaques predated most of the seeds of S. myrtacea. Dispersal success of S. myrtacea is low both by birds and macaques, thus the early fruiting by S. myrtacea does not seem to be an adaptation to maximize dispersal success by depending on resident dispersers or by avoiding intense competition for dispersers.

Dietary adaptations of temperate primates : comparisons of Japanese and Barbary macaques

Habitat, diet and leaf chemistry are compared between Japanese and Barbary macaques to reveal the similarities and differences in dietary adaptations of temperate primates living at the eastern and western extremes of the genus Macaca. Tree species diversity and proportion of fleshy-fruited species are much higher in Japan than in North Africa. Both species spend considerable annual feeding time on leaves. Japanese macaques prefer fruits and seeds over leaves, and Barbary macaques prefer seeds. These characteristics are adaptive in temperate regions where fruit availability varies considerably with season, since animals can survive during the lean period by relying on leaf and other vegetative foods. The two species are different with respect to the higher consumption of herbs by Barbary macaques, and the leaves consumed contain high condensed and hydrolysable tannin for Barbary but not for Japanese macaques. Barbary macaques supplement less diverse tree foods with herbs. Because of the low species diversity and high tannin content of the dominant tree species, Barbary macaques may have developed the capacity to cope with tannin. This supports the idea that digestion of leaves is indispensable to survive in temperate regions where fruit and seed foods are not available for a prolonged period during each year.

Coming down from the trees: Is terrestrial activity in Bornean orangutans natural or disturbance driven?

The orangutan is the worlds 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 orangutans 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.