Quantifying the direct and indirect effects of sika deer (Cervus nippon) on the prevalence of infection with Rickettsia in questing Haemaphysalis megaspinosa: A field experimental study.

Sika deer (Cervus nippon) are important hosts for all life stages of Haemaphysalis megaspinosa, a suspected Rickettsia vector. Because some Rickettsia are unlikely to be amplified by deer in Japan, the presence of deer may decrease the prevalence of Rickettsia infection in questing H. megaspinosa. As sika deer decrease vegetation cover and height and thereby indirectly cause changes in the abundance of other hosts, including reservoirs of Rickettsia, the prevalence of Rickettsia infection in questing ticks can also change. We investigated these possible effects of deer on the prevalence of infection with Rickettsia in questing ticks in a field experiment in which deer density was manipulated at three fenced sites: a deer enclosure (Deer-enclosed site); a deer enclosure where deer had been present until 2015 and only indirect effects remained (Indirect effect site); and a deer exclosure in place since 2004 (Deer-exclosed site). Density of questing nymphs and the prevalence of infection with Rickettsia sp. 1 in questing nymphs at each site were compared from 2018 to 2020. The nymph density at the Deer-exclosed site did not significantly differ from that at the Indirect effect site, suggesting that the deer herbivory did not affect the nymph density by reducing vegetation and increasing the abundance of other host mammals. However, the prevalence of infection with Rickettsia sp. 1 in questing nymphs was higher at the Deer-exclosed site than at the Deer-enclosed site, possibly because ticks utilized alternative hosts when deer were absent. The difference in Rickettsia sp. 1 prevalence between the Indirect effect and Deer-exclosed sites was comparable to that between the Indirect effect and Deer-enclosed sites, indicating that the indirect effects of deer were as strong as the direct effects. Examining the indirect effects of ecosystem engineers in the study of tick-borne diseases may be more important than previously recognized.

A simple method for calculating minimum estimates of previous population sizes of wildlife from hunting records.

Hunting records have proven useful for examining the historical status of wildlife populations. The number of animals harvested can provide information on past population sizes that would have been required to support harvest yields. Therefore, when statistical data on annual harvests are available, a minimum estimate of past population sizes can be calculated. A very simple method for estimating the sizes of historic wildlife populations using only annual hunting records and the maximum annual population increase rate is presented in this study. This method was applied to estimate past population sizes for Japanese sika deer (Cervus nippon yesoensis) in Hokkaido Island, Japan, using hunting records from 1873 to 1882, and assuming 15% and 35% population increase rates. The annual number of deer harvested during 1873 to 1882 ranged from 15,000 to 129,000. The minimum population size in 1873 was estimated as 349,000-473,000. This method was validated by applying it to the eastern population of Hokkaido Island in 1993 when the population size was approximately 260,000, and population sizes estimated by this method were 0.50-1.17 times the nominal population size. Thus, the population estimates from this method were approximately equal to or less than the expected population sizes, and this method can be used to obtain minimum estimates of wildlife populations. Because shorter durations of hunting records result in population size underestimates, it would be better to use hunting record of 10 years or longer in this method. In addition, the degree of underestimation may change with hunting pressure intensity on the populations, other causes of mortality, and maximum annual increase rates of the species. The method can be applied to any wildlife species for which records of annual harvest and maximum annual population increase rates of the species are available. The estimates obtained can provide benchmarks for the population size required for ecosystem conservation, and can be useful for wildlife management as they indicate the lowest limit to maintain the population.

Effective spatial scales for evaluating environmental determinants of population density in Yakushima macaques.

Population densities of wildlife species tend to be correlated with resource productivity of habitats. However, wildlife density has been greatly modified by increasing human influences. For effective conservation, we must first identify the significant factors that affect wildlife density, and then determine the extent of the areas in which the factors should be managed. Here, we propose a protocol that accomplishes these two tasks. The main threats to wildlife are thought to be habitat alteration and hunting, with increases in alien carnivores being a concern that has arisen recently. Here, we examined the effect of these anthropogenic disturbances, as well as natural factors, on the local density of Yakushima macaques (Macaca fuscata yakui). We surveyed macaque densities at 30 sites across their habitat using data from 403 automatic cameras. We quantified the effect of natural vegetation (broad-leaved forest, mixed coniferous/broad-leaved forest, etc.), altered vegetation (forestry area and agricultural land), hunting pressure, and density of feral domestic dogs (Canis familiaris). The effect of each vegetation type was analyzed at numerous spatial scales (between 150 and 3,600-m radii from the camera locations) to determine the best scale for explaining macaque density (effective spatial scale). A model-selection procedure (generalized linear mixed model) was used to detect significant factors affecting macaque density. We detected that the most effective spatial scale was 400 m in radius, a scale that corresponded to group range size of the macaques. At this scale, the amount of broad-leaved forest was selected as a positive factor, whereas mixed forest and forestry area were selected as negative factors for macaque density. This study demonstrated the importance of the simultaneous evaluation of all possible factors of wildlife population density at the appropriate spatial scale.

Ecosystem impacts of folivory and frugivory by Japanese macaques in two temperate forests in Yakushima.

Comparing animal consumption to plant primary production provides a means of assessing an animal's impact on the ecosystem and an evaluation of resource limitation. Here, we compared annual fruit and leaf consumption by Japanese macaques (Macaca fuscata) relative to the annual production of these foods in the lowlands and highlands of Yakushima Island, Japan. We estimated consumption by macaques by the direct observation of macaque groups for 1 year in each habitat. We estimated leaf production as the sum of leaf litter fall (corrected for the effect of translocated organic and inorganic matter) and folivory by insects (assumed to be 10%) and by macaques. We estimated fruit production as the sum of fruit litter fall and consumption by birds (estimated by the seed fall) and macaques. The impact of macaque folivory at the community level was negligible relative to production (∼0.04%) compared with folivory by insects (assumed to be 10%); however, for some species, macaque folivory reached up to 10.1% of production. Tree species on which macaques fed did not decline in abundance over 13 years, suggesting that their folivory did not influence tree species dynamics. For the three major fleshy-fruited species in the highland site, macaques consumed a considerable portion of total fruit production (6-40%), rivaling the consumption by birds (32-75%). We conclude that at the community level, macaque folivory was negligible compared with the leaf production, but frugivory was not.

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.

Acoustic identification of eight species of bat (mammalia: chiroptera) inhabiting forests of southern hokkaido, Japan: potential for conservation monitoring.

Assessing the impact of forest management on bat communities requires a reliable method for measuring patterns of habitat use by individual species. A measure of activity can be obtained by monitoring echolocation calls, but identification of species is not always straightforward. We assess the feasibility of using analysis of time-expanded echolocation calls to identify free-flying bats in the Tomakomai Experimental Forest of Hokkaido University, Hokkaido, northern Japan. Echolocation calls of eight bat species were recorded in one or more of three conditions: from hand-released individuals, from bats flying in a confined space and from bats emerging from their roost. Sonograms of 171 calls from 8 bat species were analyzed. These calls could be categorized into 3 types according to their structure: FM/CF/FM type (Rhinolophus ferrumequinum), FM types (Murina leucogaster, Murina ussuriensis, Myotis macrodactylus and Myotis ikonnikovi) and FM/QCF types (Eptesicus nilssonii, Vespertilio superans and Nyctalus aviator). Sonograms of the calls of R. ferrumequinum could easily be distinguished from those of all other species by eye. For the remaining calls, seven parameters (measures of frequency, duration and inter-call interval) were examined using discriminant function analysis, and 92% of calls were correctly classified to species. For each species, at least 80% of calls were correctly classified. We conclude that analysis of echolocation calls is a viable method for distinguishing between species of bats in the Tomakomai Experimental Forest, and that this approach could be applied to examine species differences in patterns of habitat-use within the forest.

Altitudinal and seasonal variations in the diet of Japanese macaques in Yakushima.

Altitudinal and seasonal variations in the diet of Japanese macaques in Yakushima, southwestern Japan, were studied for 2 years by means of fecal analysis. The altitudinal range of fecal samples collected was 30 m to 1,203 m above sea level, and it was divided into three zones: low-zone forest (0-399 m), middle-zone forest (400-799 m), and high-zone forest (800 m-1,230 m). There was a considerable altitudinal and seasonal variation in the macaques' diet. Seed/fruit and animal matter were eaten more in the lower zones, whereas more fiber and fungi were consumed in the higher zones. In all of the zones, they ate seed/fruits the most in autumn (September-November) and the least in spring (March-April). They ate fibrous food the most in spring and the least in autumn. Macaques relied on seed/fruits heavily in the lower zone for a longer period than in the higher zones. Macaques in the high-zone forest ate almost no seed/fruit foods from March to May. Altitudinal variations in availability of seed/fruit foods seem to have influenced the altitudinal variations in diet. Total basal area of seed/fruit-food trees, species richness of seed/fruit-foods, main seed/fruit-food types available, and annual fleshy-fruit production all decreased with increasing altitude. Both interannual variation and annual cyclicity of diet were found in all zones.

Supra-annual variation in the influence of Myrica rubra fruit on the behavior of a troop of Japanese macaques in Yakushima.

Observations were made on a well-habituated natural troop of Japanese macaques (Macaca fuscata yakui), living in warm-temperate, lowland forest in Yakushima. Between mid-May and the end of June the macaques feed on the fruit of the evergreen tree Myrica rubra (Myricaceae). The fruit of this species are abundant in some years and scarce in others. Data were compared for two heavy-fruiting years (1988 and 1990) and one poor-fruiting year (1991) to examine the influence of fruit availability on patterns of foraging, ranging, and the frequency of inter-troop encounters. In both heavy-fruiting years M. rubra fruit accounted for a maximum of over 70% of foraging time, compared with a maximum of <5% in the poor-fruiting year. Heavy fruiting was also associated with a marked decrease in the overall time spent foraging. In early May of all three years troop movements were largely confined to northern parts of the home range. By early June of both heavy-fruiting years ranging had shifted to the south-west, and included an area with a high concentration of M. rubra trees. This area was rarely visited at other times, and was not visited during the study period in the poor-fruiting year. The overlap in range-use between the two heavy-fruiting years was significantly greater than that between the heavy-fruiting years and the poor-fruiting year. Heavy fruiting was also associated with an increase in the frequency of inter-troop encounters. © 1995 Wiley-Liss, Inc.