Summer dietary compositions of sika deer on Yakushima Island,southern Japan

Summer food habits of Sika deer (Cervus nippon yakushimae) on Yakushima Island, southern Japan, were studied focusing in particular onPseudosasa owatarii, a dwarf bamboo native to the island. Fecal analyses showed that the food habit of the Sika deer on Yakushima island was plastic: the dietary composition was dominated by graminoids (ca. 90%), particularlyP. owatarii (50%–60%) in the bamboo grassland of theP. owatarii zone above the timber line, whereas leaves and non-leafy parts of woody plants were important in the crytomeria forest and evergreen broad-leaved forest below the timber line. The maximum culm age ofP. owatarii was 8.5 yr, but the majority of leaves (97.8%) were younger than 2.5 yr. Since the weight contribution of the leaves is great (27.7%) and the turnover rate is high, theP. owatarii grassland affords a good foraging place for the Sika deer on Yakushima Island.     

Food habits of sika deer in the Shiranuka Hills,eastern Hokkaido: a northern example from the north–south variations in food habits in sika deer

Seasonal changes in the composition of the diet of the sika deer population in the Shiranuka Hills, eastern Hokkaido, in 1998 were determined by fecal analyses. The deer were dependent on Sasa nipponica, a dwarf bamboo, throughout the year, particularly in winter when it accounted for as much as 77.7% of the diet. It accounted for 33.1% and 45.6% in spring and summer, respectively, and this decreased to 12.2% in autumn. Besides S. nipponica, all the graminoid categories accounted for large amounts (66–96.7%), while dicotyledonous plants accounted for little (3–8%) except in autumn when they accounted for 31%. The strong dependence of the Shiranuka population on graminoids was different from other Hokkaido deer populations, for example the population from Ashoro/Onbetsu and the extremely high density population on Nakanoshima Island. In spite of these differences, food for all Hokkaido sika deer was poor in winter. Along the north–south geographical cline in the food composition of sika deer along the Japanese archipelago, the Shiranuka population was positioned as a grazer type, in contrast to the southern populations. However, it is important to note that variations are great among local populations in Hokkaido.     

Effects of deer,mice and dwarf bamboo on the emergence,survival and growth of <Emphasis Type="Italic">Abies homolepis</Emphasis> (Piceaceae) seedlings

In the subalpine mixed forest of Mt. Ôdaigahara, mid-western Japan, the understory is dominated by dwarf bamboo (Sasa nipponica), which is the major forage of overly populous sika deer (Cervus nippon). In the present study, we monitored the survival and growth of Abies homolepis seedlings over 5years to determine how they responded to the experimental removal of dwarf bamboo and to the exclusion of sika deer and mice (Apodemus argenteus and A.speciosus). Deer and dwarf bamboo reduced the survival of seedlings but had different effects on growth. The stems of seedlings were shorter in the presence of deer, indicating that taller seedlings were apt to be browsed by deer, whereas the diameters of seedlings were smaller in the presence of dwarf bamboo, mainly owing to its shading effect. The presence of mice decreased the number of seedlings germinating in a particular site, but had no effect on seedling survival after germination. There was no significant indirect effect whereby the survival of seedlings was predicted to be facilitated by the decreased biomass of bamboo because of grazing by deer. We supposed that this might be because the direct negative effect of deer was so large as to conceal the positive indirect effect.     

How do deer affect tree seedlings on a dwarf bamboo-dominated forest floor?

A field experiment was conducted in Mt. Ôdaigahara, west-central Japan, to reveal the effects of two herbivores, Sika deer (Cervus nippon) and mice (Apodemus spp.), on the seedlings of five cohorts of three tree species, Abies homolepis, Fraxinus lanuginosa f. serrata and Fagus crenata. The forest floor of the study site was covered with dwarf bamboo, Sasa nipponica, which would also affect the seedlings. Eight combinations of three treatments were set: exclusion of deer, exclusion of mice and removal of dwarf bamboo. Deer were expected to affect the seedlings not only negatively by foraging but also positively by browsing dwarf bamboo that overshadows seedlings. The survival of these cohorts was analyzed by survival analysis and the differences in their survival between the treatments were investigated. The results of the experiment showed that: (1) exclusion of deer increased the aboveground biomass of dwarf bamboo and made it more difficult for seedlings to survive under the regenerated dwarf bamboo stand, and (2) deer negatively affected the seedlings, but they had positive, indirect effects in some cohorts by decreasing the aboveground biomass of dwarf bamboo. However, such effects were not detected in some cohorts which were affected excessively by deer or dwarf bamboo. No clear results on the effect of mice on seedlings were obtained. We conclude that a positive, indirect effect of deer on seedling survivorship would be observable when the magnitude of a positive, indirect effect caused by decreasing dwarf bamboo is larger than that of a negative, direct effect of deer.     

Characteristics of Sasa nipponica grassland as a summer forage resource for sika deer on Mt Ohdaigahara, central Japan

Recently, the sika deer, Cervus nippon Temminck, population has increased on Mt Ohdaigahara, central Japan. The dwarf bamboo, Sasa nipponica Makino et Shibata, is a primary forage plant for sika deer in this area. To demonstrate the characteristics of S. nipponica grassland, especially as summer forage for sika deer, the habitat use intensity of sika deer was estimated by fecal densities, and biomass, growth rate, removal by deer and crude protein content were examined. Sika deer utilized the S.nipponica grassland on Mt Ohdaigahara during summer when the biomass, growth rate and crude protein content of S. nipponica were high. The recent increase in the deer population seems to be partly due to S.nipponica grassland being a favorable summer habitat.     

Relationship between a high density of sika deer and productivity of the short-grass (<Emphasis Type="Italic">Zoysia japonica</Emphasis>) community: a case study on Kinkazan Island,northern Japan

An extremely high-density (ca. 800 deer km^–2) wild sika deer population uses a short-grass community dominated by Zoysia japonica on Kinkazan Island in northeastern Japan. To explain why the density of wild deer is quite high on the Zoysia community, (1) we quantified the seasonal productivity of the Zoysia community, (2) we compared food availabilities among the plant communities, and (3) we described the habitat selection by the deer in different seasons. Food availability was greater on the Zoysia community than in the forest understory from spring to fall. The productivity of the Zoysia community was high enough to support the actual high density of the deer (814 deer km^–2) in summer. However, the productivity markedly decreased in winter, when the deer density decreased to less than half (358 deer km^–2) of the summer value. In contrast, the deer density of the adjacent forests was highest in winter (154 deer km^–2) and lowest in spring (19 deer km^–2). These results suggest that the deer using the Zoysia community in summer left and were absorbed into the adjacent forest in winter. If such an adjacent community were absent, many deer would not survive, and consequently the deer density on the Zoysia community in summer would not be so high. This intercommunity movement is particularly important for the deer using a plant community like the Zoysia community, which is highly productive but has a small standing biomass.     

A case study on the effects of a transmission-line corridor on Sika deer habitat use at the foothills of Mt Goyo,northern Honshu,Japan

Forage biomass and habitat use of Sika deer (Cervus nippon) at a transmission-line corridor were studied at the foothills of Mt Goyo, northern Japan. Summer forage biomass in the corridor was five times greater than in the adjacentBetula grossa forest. Among the plants that increased in the corridor,Sasa nipponica (a dwarf bamboo), an important forage plant for Sika deer, was predominant. Winter utilization ofS. nipponica by Sika deer was slightly heavier in the corridor, and estimated removal ofSasa leaves was twice as great there as in the forest. However fecal pellets were more prevalent in the adjacent forest in winter. Sika deer seemed to use the transmission-line corridor as a feeding site and the adjacent forest as cover as it reduces wind speed. A transmission-line corridor is more beneficial than a large clear-cut area because it provides more forest edges.     

Food habits of sika deer and nutritional value of sika deer diets in eastern Hokkaido, Japan

Rumen content analysis and field observations were used to investigate the food habits and diet quality of sika deer (Cervus nippon yesoensis Heude) from 1991 to 1993 in eastern Hokkaido, Japan. Diets varied seasonally, with deer consuming graminoids and browse in winter, forbs and agricultural crops in spring and summer and all of these plant foods in autumn. Eighty-four plant species with sika deer bite marks were identified and their use also varied seasonally. The diversity of food resources available provided both critical protein and digestible energy, allowing for physiological maintenance and seasonal growth. With these high-quality diets, deer maintained good body condition in eastern Hokkaido, where the population density was relatively low.     

Edge effects created by clear-cutting on habitat use by sika deer on Mt. Goyo,northern Honshu,Japan

Clear-cutting increased the species diversity and amount of undergrowth plants in a habitat of Sika deer (Cervus nippon) on Mt. Goyo, northern Japan. The number of species increased from 15 to 48 as a result of clear-cutting. Among the plants,Sasa nipponica (a dwarf bamboo), an important forage plant for Sika deer, was predominant. Fecal pellets of deer were abundant in the forest and at the “adjacent zone” (from the edge to 150 m out of the forest) and thereafter decreased suddenly. The intensity of utilization ofSasa nipponica was also heavy in the forest, moderate at the adjacent zone and light 200 m from the forest edge. Since the amount of the bamboo in the forest was small, the removal of bamboo was greatest at the adjacent zone. Clear-cutting creates a favorable feeding area for Sika deer in this zone by increasing the available plant production and securing forest cover.     

Dwarf bamboo as an ecological filter for forest regeneration

Quantitative relationships between the aboveground biomass of dwarf bamboo, Sasa nipponica, and the survivorship and emergence of seedlings of Abies homolepis, Fraxinus lanuginosa f. serrata, and Fagus crenata were estimated. We show that dwarf bamboo acts as an ecological filter since the responses of tree species differ according to the biomass of dwarf bamboo. Deer exclusion without management of dwarf bamboo would make it impossible for any tree species to regenerate due to rapid increases in the biomass of dwarf bamboo.     

Food habits of Sika deer on Mt. Goyo,northern Honshu

The food habits of Sika deer (Cervus nippon) on Mt. Goyo, northern Honshu (the main island of Japan), were studied. Among 88 plant species found in surveys, 36 species (40.9%) were eaten by the deer. The percentage frequency of feeding (F) was highest for graminoids (59.6%), followed by browses (16.6%) and ferns (7.7%) and lowest for forbs (5.4%). Though various kinds of plants were eaten, only limited species formed the bulk of the deer dietary composition.Sasa nipponica, in particular, was the most important constituent, ranking highest inF value (80.3%) and being the dominant component of feces. Fecal analyses revealed that, 1) in areas lower than 900 m in altitude,Sasa nipponica accounted for between 20 and 60% of the content and that pasture grasses were also important, and 2)S. nipponica occupied more than 50% in montane areas higher than 900 m, and even more than 80% in winter. The utilization ofS. nipponica as a forage for Sika deer was discussed. This plant can be a key forage species because it meets all the necessary conditions: abundance and stability of supply, palatability, nutritive quality and tolerance to grazing.     

Diet of Eurasian lynx, Lynx lynx, in the boreal forest of southeastern Norway: the relative importance of livestock and hares at low roe deer density

The year-round food habits of lynx were studied using radio-telemetry and snow-tracking in the boreal forest of southeastern Norway. The main objectives of the study were to clarify the importance of domestic sheep and small prey species in the diet of lynx in an area with a very low density of roe deer. During the period 1995–1999, we found 193 scats and 358 kills made by lynx. Our results indicate that roe deer were the most common prey species (contributing to 83 and 34% of the biomass consumed in winter and summer, respectively), although a wide range of other species were also found, including mountain hares, tetranoids, red foxes, domestic sheep, wild reindeer, and even moose. Most of the diet was obtained by predation, although we did document several cases of scavenging. Roe deer were more important in the diet in winter than in summer, perhaps because they were easier to locate in winter as they clustered around feeding sites. In summer, domestic sheep and small prey increased in importance. Despite the very low density of roe deer in this study area, lynx seemed to still specialise on them, although domestic sheep did constitute a significant amount to their diet, especially for males and yearlings. However, the contribution of sheep to summer diet was far from that expected if their relative density was considered.     

Seasonal migration patterns of female sika deer in eastern Hokkaido,Japan

Fifty-seven female sika deer (Cervus nippon yesoensis), captured at the wintering area in the Shiranuka Hills in eastern Hokkaido, Japan, were radio-tracked during 1997–2001 to examine the factors affecting seasonal migration at the individual-landscape level. Ten of the 57 deers migrated between low-altitude summer home ranges and intermediate-altitude winter home ranges (the upward migrants). Twenty-nine migrated between high-altitude summer home ranges and intermediate-altitude winter home ranges (the downward migrants). Twelve used the intermediate-altitude home ranges all year round (the non-migrants). The remaining six were unknown. The summer home ranges of deer were widely scattered over an area of 5734km². Migration distances ranged between 7.2 and 101.7km. Deer showed high site fidelities to their seasonal home ranges. The upward migrants wintered in areas of less snow, higher quality of bamboo grass, and more coniferous cover than their summer home ranges. The downward migrants wintered in areas of less snow, higher quality of bamboo grass, higher winter temperature, and more southern slopes, but less coniferous cover than their summer home ranges. The non-migrants used year-round ranges with little snow, high quality of bamboo grass, and sufficient coniferous cover. We suggest that snow cover and bamboo grass are the factors affecting seasonal migration of the population and that coniferous cover is another factor for the upward migration.     

Dry season diets of sympatric ungulates in lowland Nepal: competition and facilitation in alluvial tall grasslands

Based on microhistological analyses of faecal material, we compared the early dry season diets of greater one-horned rhinoceros Rhinoceros unicornis, swamp deer Cervus duvauceli and hog deer Axis porcinus, which inhabit the same alluvial grassland habitat complex in lowland Nepal. Their diets were quite similar, both at the forage category level and within subcategories of graminoids and woody plants. Early successional tall grasses, especially Saccharum spontaneum, were the dominant food of all three species, underlining the key role of the threatened alluvial floodplains in large mammal conservation in South Asia. The two deer species ate significantly more graminoids (>66.5%) than did rhino (45.5%), and although they did not differ in proportions of graminoids, swamp deer ate significantly more late successional tall grasses (Narenga porphyrocoma and Themeda spp.) and short grasses (mainly Imperata cylindrica) than hog deer. The two deer consumed almost equal proportions of woody browse (ca. 10%), significantly less than that of rhino (33.0%). The prediction of the Jarman–Bell hypothesis, that large-bodied herbivores are less selective and subsist on lower quality graminoids than smaller ruminants, was not supported by the data. Based on this and previous studies in the same area we propose a conceptual model where the larger megaherbivores (rhino and elephant Elephas maximus) facilitate the smaller swamp deer and hog deer during the monsoonal growing season, while the smaller and more selective deer species outcompete the larger during the dry season when food is more limited. Owing to the all-year sprouting ability of S. spontaneum, facilitation may occur also in the dry season with low numbers of megaherbivores, thus accentuating competitive exclusion at higher deer densities.     

Food habits of sambar <Emphasis Type="Italic">Cervus unicolor</Emphasis> at the Horton Plains National Park,Sri Lanka

The sambar Cervus unicolor has a large population in southern Asia. In Sri Lanka, its habitat use is most common among the four cervid species. We carried out an analysis of the food habits of sambar in the Horton Plains National Park from January to November 1991 by using the quantitative fecal analysis method. Analyses of sambar feces collected at the three habitats (two grassland communities and a forest) in the Horton Plains National Park found that the sambar diet largely consisted of graminoid leaves; 45–78% in grasslands and 43–60% in the forest. The analyses also found that the proportion of graminoid leaves in early summer (52–78%) was greater than it was in winter (43–49%). In spring and summer, different varieties of graminoids were consumed by sambar, whereas dicotyledonous plants were consumed in winter. The fecal nitrogen level did not fall below the critical level of their nutrition (1.1%), indicating that sambar food quality in the Horton Plains was sufficient. A large proportion of Pennisetum spp., exotic grasses (10–39%) were found in the fecal compositions collected in the Pennisetum community. This could be because of the high biomass and high nitrogen concentration (approximately 2.4%) of Pennisetum and its low-growing nature. From the present study, we found that in order for sambar, a native ungulate, not to increase in number, the growth of exotic plants such as Pennisetum needs to be controlled.     

Age-sex differences in the diets of Sika deer on Kinkazan Island,northern Japan

Dietary differences in the ages and sexes of Sika deer (Cervus nippon) were studied on Kinkazan Island, northern Japan from October 1990 to July 1991. Larger deer consumed more graminoids than smaller deer except in spring, and less dicotyledonous leaves in all the seasons. Fecal nitrogen concentrations were lower in larger deer than in smaller deer in all the seasons. The age-sex differences in foods were smaller in summer and winter when foods were most abundant and scarce, respectively, while they were greater in spring and autumn when food availability was intermediate.     

Structure and dynamics of Japanese beech (Fagus japonica Maxim.) stools and sprouts in the regeneration of the natural forests

Structure and spatial distribution of stools and root-collar sprouts of Japanese beech (Fagus japonica) were studied to clarify the regeneration processes of the stool and the population, and the ecological importance of this stool formation in five quadrats of the natural forests with different forest floor vegetation on the Pacific side of Japan. F. japonica dominates in the canopy of each quadrat. Most of sprouts formed a circle around the root-collar and lowest parts of the parent stems of the stool with the youngest sprouts at the periphery. The regeneration by seedlings was slight especially on the forest floor vegetation of the dwarf bamboo Sasa. The variety of size structure of stems and the existence of dead traces and/or dead center in each stool, the apparent difference in stool size, and the aggregations of stools in the forests suggest that stool expansion and long persistence of the stool at a given location may contribute to compensate for the scarcity of regeneration by seedlings inhibited by dwarf bamboo, and by the other shrubs and herbs.     

Regeneration of beech (Fagus crenata) after the simultaneous death of undergrowing dwarf bamboo (Sasa kurilensis)

Seedling recruitment and survivorship of beech (Fagus crenata) were studied with special reference to the simultaneous death of undergrowing bamboo (Sasa kurilensis). The survival rate of beech seedlings on the floor whereSasa had withered was much higher than that on the floor whereSasa survived. Damping off caused the largest mortality among beech seedlings. However, the allocation pattern of matter to different parts of the seedlings indicated that their survival was greatly affected by production economy. The dense cover of dwarf bamboo prevented the establishment of beech seedling banks on the forest floor. The interval between the times when simultaneous death ofSasa occur and the length of its recovery period are thus important factors controlling the dynamics of beech forests in Japan.     

Botanical and chemical composition of rumen contents of Sika deer on Mt Goyo,northern Japan

Botanical and chemical compositions of the rumen contents of 58 Sika deer on Mt Goyo, northern Japan, collected from summer of 1988 to spring of 1989, were analyzed.Sasa nipponica, a dwarf bamboo, was important in summer (35.0%) and winter (61.4%), but it decreased to 5.6% and was replaced by browse leaves in fall, and to 28.0% and was replaced by dead leaves, twigs and bark of woody plants in winter. Crude protein was 20–25% lower in the washed fraction than in the gross fraction. It was highest (16.2%) in summer and lowest (8.6%) in winter. High protein content in summer and fall foods was attributed to forb and browse leaves. Seasonal fluctuation in protein content in the foods of these Sika deer was greater than red deer on Rhum, Scotland and smaller than wapiti in Yellowstone National Park, Wyoming, USA. Climatically, winter severity of Mt Goyo is intermediate between the two localities, which seems to explain the seasonal fluctuation of the protein level. Crude fiber wasca 33% in the ‘washed’ fraction, and did not change seasonally.     

Effects of dwarf bamboo (Fargesia denudata) density on biomass, carbon and nutrient distribution pattern

Dense dwarf bamboo population is a structurally and functionally important component in many subalpine forest systems. To characterize the effects of stem density on biomass, carbon and majority nutrients (N, P, K, Ca and Mg) distribution pattern, three dwarf bamboo (Fargesia denudata) populations with different stem densities (Dh with 220 ± 11 stems m^?2, Dm with 140 ± 7 stems m^?2, and Dl with 80 ± 4 stems m^?2, respectively) were selected beneath a bamboo-fir (Picea purpurea) forest in Wanglang National Nature Reserve, Sichuan, China. Leaf, branch, rhizome, root and total biomass of dwarf bamboo increased with the increase of stem density, while carbon and nutrient concentrations in bamboo components decreased. Percentages of below-ground biomass and element stocks to total biomass and stocks decreased with the increase of stem density, whereas above-ground biomass and element stocks exhibited the opposite tendency. Moreover, more above-ground biomass and elements were allocated to higher part in the higher density population. In addition, percentages of culm biomass, above-ground biomass and element stocks below 100 cm culm height (H₁₀₀) increased with the increase of stem density, while percentages of branch and leaf biomass below H₁₀₀ decreased. Pearson’s correlation analyses revealed that root biomass, above-ground biomass, below-ground biomass and total biomass significantly correlated to leaf biomass in H_100?200 and total leaf biomass within high density population, while they significantly correlated to leaf biomass in H_50?150 within low density population. The results suggested that dwarf bamboo performed an efficient adaptive strategy to favor limited resources by altering biomass, carbon and nutrients distribution pattern in the dense population.