Tsutomu Enoki

Microtopography and distribution of canopy trees in a subtropical evergreen broad-leaved forest in the northern part of Okinawa Island, Japan

The spatial distribution of trees in relation to topography was investigated using two topographic indices, slope steepness and slope configuration, in a 4-ha plot in a subtropical evergreen forest in the northern part of Okinawa Island, Japan. Most species showed an aggregated distribution and patch size was related to various microtopographical features, including small ridges and valleys. In a cluster analysis based on the dissimilarity of each species’ distribution, the species were divided into three major clusters. A significant correlation between the dissimilarity and the distance between each species on the axis of the two topographic indices indicated that species association was, in part, explained by the topographic indices. I suggest that species distributed on steep and concave slopes regenerate depending on disturbances such as landslides on unstable topography, whereas species distributed on ridges and upper slopes regenerate depending on the canopy gap. A number of species that were less abundant in the 4-ha plot occurred in the riparian area near a stream, where the density of more abundant species was low. The results of this study support the existence of habitat niche divergence related to topography in a subtropical evergreen broad-leaved forest.

Topographic variations of soil properties and stand structure in a Pinus thunbergii plantation

Soil properties and above-and belowground forest structure were studied across various topographies in a 20-year-oldPinus thunbergii Parl. plantation on Mt Tanakami, Japan. The soil properties and stand structure varied greatly with slope position fromridge top to valley floor. Soil thickness, fine soil content and soil moisture content were greater in lower slope positions. The amount of organic carbon in the forest floor was greater in upper slope positions. The organic carbon content in the mineral soil was slightly greater in lower slope positions. These changes in soil properties suggested an upslope decrease in decomposition rate and water and/or nutrient availability. The aboveground structure ofP. thunbergii was more developed at lower slope positions. The mean stem diameter, height and volume ofP. thunbergii increased downslope with decreasing tree density. However, fine root biomass increased greatly upslope. This inverse relationship between tree height and fine root biomass indicated morphological plasticity ofP. thunbergii in exploiting environmental heterogeneity. Variations in soil-plant interactions in the stand along various topographies caused spatial heterogeneity in the accumulation pattern of organic matter in plants and the soil.

Saplings distribution in relation to topography and canopy openness in an evergreen broad-leaved forest

We examined the spatial distribution of saplings in relation to topography and stand structure in a subtropical evergreen broad-leaved forest in the northern part of Okinawa Island, Japan. The distributions of most species were influenced mainly by topography but also by canopy openness. Species were arranged along a topographical gradient from concave areas with low canopy openness to convex areas with high canopy openness. Canopy species were arranged along a gradient from unstable fertile areas to stable infertile areas. Species occurring mostly in convex areas had a narrower distribution range than those in concave sites. Thus, habitat heterogeneity provided by topography and stand structure appears to be important factors for maintaining high species diversity in the understory of this subtropical evergreen broad-leaved forest.

Leaf litter decomposition and nutrient dynamics in a subtropical forest after typhoon disturbance

Decomposition of typhoon-generated and normal leaf litter and their release patterns for eight nutrient elements were investigated over 3 yr using the litterbag technique in a subtropical evergreen broad-leaved forest on Okinawa Island, Japan. Two common tree species, Castanopsis sieboldii and Schima wallichii, representative of the vegetation and differing in their foliar traits, were selected. The elements analyzed were N, P, K, Ca, Mg, Na, Al, Fe and Mn. Dry mass loss at the end of study varied in the order: typhoon green leaves > typhoon yellow leaves > normal leaves falling for both species. For the same litter type, Schima decomposed faster than Castanopsis. Dry mass remaining after 2 yr of decomposition was positively correlated with initial C:N and C:P ratios. There was a wide range in patterns of nutrient concentration, from a net accumulation to a rapid loss in decomposition. Leaf litter generated by typhoons decomposed more rapidly than did the normal litter, with rapid losses for N and P. Analysis of initial quality for the different litter types showed that the C:P ratios were extremely high (range 896 − 2467) but the P:N ratios were < 0.05 (range 0.02 − 0.04), indicating a likely P-limitation for this forest. On average 32% less N and 60% less P was retranslocated from the typhoon-generated green leaves than from the normal litter for the two species, Castanopsis and Schima. An estimated 2.13 g m–2 yr–1 more N and 0.07 g m–2 yr–1 more P was transferred to the soil as result of typhoon disturbances, which were as high as 52% of N and 74% of P inputted from leaf litter annually in a normal year. Typhoon-driven maintenance of rapid P cycling appears to be an important mechanism by which growth of this Okinawan subtropical forest is maintained.

Nutrient-uptake and nutrient-use efficiency ofPinus thunbergii Parl. along a topographical gradient of soil nutrient availability

To examine responses of a plant species to nutrient availability, we investigated changes in soil nutrient availability, litterfall production and nutrient content in litterfall along a topographic gradient in aPinus thunbergii Parl. plantation. Responses were evaluated in terms of three efficiency indices: (i) nutrient-uptake efficiency (the ratio of nutrient return in litterfall to soil nutrient availability); (ii) nutrient-use efficiency (the ratio of litterfall mass to nutrient return in litterfall); and (iii) nutrient-response efficiency (the ratio of litterfall mass to soil nutrient availability). These indices can distinguish the ability of a species to acquire nutrients and its ability to use them in litterfall production. Nitrogen and phosphorus availabilities in soil were lower in upper slope positions. The three efficiencies were higher in upper slope positions and negatively correlated with soil nutrient availability for both nitrogen and phosphorus. An increase in nutrient-response efficiency was achieved by both increases in nutrient-uptake and nutrient-use efficiencies.

Nitrogen resorption from needles of Pinus thunbergii Parl. growing along a topographic gradient of soil nutrient availability

To examine the relative importance of nutrient resorption in increasing the nutrient-use efficiency of Pinus thunbergii Parl., we investigated the nitrogen contents of green and senescent needles of P. thunbergii trees growing at five positions along a slope (LS, lower slope; TR, transitional site; MS, middle slope; US, upper slope; RG, ridge) and found that soil nitrogen availability tended to decrease upslope. Nitrogen concentration in green and senescent needles decreased upslope. Nitrogen resorption efficiencies (percentage change in nitrogen content between green and senescent needles) increased upslope from 43 to 77% with decreasing soil nitrogen availability. Nitrogen resorption efficiency was related to green needle dry mass per unit length, but there was no clear correlation between nitrogen resorption efficiency and nitrogen content in green needles. We concluded that the increase in nitrogen resorption efficiency of P. thunbergii enhanced the nitrogen-use efficiency as a response to the low nitrogen availability.

Ecological studies on subtropical evergreen broad-leaved forest in Okinawa, Japan: Litter production and nutrient input

Patterns of litterfall and nutrient input in a subtropical evergreen broad-leaved forest in northern Okinawa, Japan, were studied during May, 1996–April, 1999. The mean annual rate of litterfall in the five sampling plots ranged from 6.84 to 8.93 Mg ha−1 yr−1, of which 63.3–68.5% were leaves; 22.4–29.1% woody parts (including branches < 5.0 cm in diameter and bark); 2.8–5.0% sexual organs and 4.6–6.3% miscellaneous material. Significant differences were found among plots and among years. Significantly monthly differences pronounced seasonal patterns in litterfall were observed. Total litterfall and leaf litter showed negative correlations with relative basal area of the dominant species,Castanopsis sieboldii; and showed positive correlations with mean height of the stands. The dominant species,C. sieboldii produced an average of 2.36 Mg ha−1 yr−1 of leaf litter, which covered 30.5% of the annual litter production, and the nutrient input from those litterfall contributed 32.3, 28.3, 30.2, 22.2, 32.5, and 30.5% of total N, P, K, Ca, Mg, and Na, respectively. Nutrient use efficiency in litter production was high, especially for P and K compared with other broad-leaved forests in Japan indicating that P and K may be limiting in Okinawan evergreen broad-leaved forest.

Initial nitrogen content and topographic moisture effects on the decomposition of pine needles

The effects of litter quality and site characteristics on the decomposition process were investigated using a litterbag method. Pine needle litters with differing nitrogen concentrations (0.8, 0.6 and 0.4%) were placed on the upper and lower slopes of a Pinus thunbergii Parl. plantation. After both 3 and 6 months, the mass of decomposing litter with the lower nitrogen concentration was larger than the litter with higher nitrogen concentrations. After 9 months, there were no significant differences in the litter mass remaining, regardless of the initial nitrogen concentration. Moisture content in the litter was always higher on the lower slope, although the mass of litter was smaller. Nitrogen concentration of the decomposing litter increased linearly with accumulated mass loss. The increase in nitrogen concentration of decomposing litter was greater on the lower slope, but this increase did not differ between initial nitrogen concentrations. The nitrogen release from the decomposing litter with higher initial nitrogen concentration was larger than the release from litter bags with lower nitrogen concentrations. This result suggests that there may be positive feedback between soil nutrient availability, litter quality and nutrient release from decomposing litter at the intraspecific level.

Effects of sika deer (Cervus nippon) and dwarf bamboo (Sasamorpha borealis) on seedling emergence and survival in cool-temperate mixed forests in the Kyushu Mountains

Effects of sika deer (Cervus nippon) and dwarf bamboo (“sasa;” Sasamorpha borealis) on seedling emergence and survival were investigated in cool-temperate mixed forests in the Kyushu mountain range, Japan. We compared the effects of sika deer between two sites with different sika deer densities. One site (no-sasa forest) has sparse cover of S. borealis and a high density of sika deer, and the other site (sasa forest) has dense cover of S. borealis with a low density of sika deer. In the no-sasa forest, more seedlings emerged and the survival rate of the seedlings was higher compared with the sasa forest. Compared with the sasa forest, the canopy in the no-sasa forest was more open, the organic layer was shallower and drier, the mean daily soil temperature was higher, and soil temperature fluctuated more widely. Those environmental parameters did not differ between unfenced and fenced plots. The comparison between the sasa and no-sasa forests suggested that removal of S. borealis by sika deer had an indirect positive effect on seedling emergence and survival. However, the fenced-plot experiments indicated that sika deer inhibited seedling emergence and survival. These contradictory conclusions could result from the much larger negative effects of S. borealis on seedling emergence and survival compared with the effects of sika deer browsing.

Determination and potential importance of diterpene (kaur-16-ene) emitted from dominant coniferous trees in Japan

Reactive volatile organic compounds (VOCs) are known to affect atmospheric chemistry. Biogenic VOCs (BVOCs) have a significant impact on regional air quality due to their large emission rates and high reactivities. Diterpenes (most particularly, kaur-16-ene) were detected in all of the 205 enclosure air samples collected over multiple seasons at two different sites from Cryptomeria japonica and Chamaecyparis obtusa trees, the dominant coniferous trees in Japan,. The emission rate of kaur-16-ene, was determined to be from 0.01 to 7.1 μg dwg(-1) h(-1) (average: 0.61 μg dwg(-1) h(-1)) employing branch enclosure measurements using adsorbent sampling followed by solid phase-liquid extraction techniques. The emission rate was comparable to that of monoterpenes, which is known major BVOC emissions, collected from the same branches. In addition, total emission of kaur-16-ene at 30°C was estimated to exceed that of total anthropogenic VOC emissions.