Shin-ichiro Aiba

Effects of topography on tropical lower montane forests under different geological conditions on Mount Kinabalu, Borneo

Species composition and forest structure change with topography.However, mechanisms for topographical vegetation changes are still not wellunderstood, because a topographical gradient is a complex environmentalgradientinclusive of many factors. The foot of Mt. Kinabalu is covered with three typesof geological substrates, i.e. Quaternary and Tertiary sedimentary rocks andultrabasic (serpentine) rock. Quaternary and Tertiary sedimentaryrocks are different in site age, but controlled in primary minerals. Tertiarysedimentary and ultrabasic rocks are contrasting in primary minerals, but arecomparable in age. This setting provides an opportunity to examine thevegetation differentiation along topographical gradients that are contrastinginmajor-nutrient supply due to the difference in site age and parent rock.We established a total of nine study plots by choosing three topographicalunits(ridge, middle- and lower-slope) on each substrate inthe tropical lower montane forest. Pool size and supply of soil N and Pdecreased upslope on each substrate, and the magnitude of the reduction fromslope to ridge decreased in the order of Quaternary sedimentary > Tertiarysedimentary > ultrabasic rock. Between-substrate difference in soilnutrient condition was greater on the lower-slopes than the ridges.Maximum tree size decreased and stem density increased upslope on eachsubstrate. Detrended correspondence analysis demonstrated that speciescomposition also changed along topographical gradients on all substrates.However, the magnitude of topographical changes in forest structure and speciescomposition varied with substrate and decreased from Quaternary sedimentary>Tertiary sedimentary > ultrabasic rock. The greatest between-substratedifference in vegetation occurred on the lower-slopes. Accordingly, ourresults suggest that the magnitude of vegetation changes due to topographybecomes smaller with decreasing pool size and supply of nutrients.

An estimate of the number of tropical tree species

Significance People are fascinated by the amazing diversity of tropical forests and will be surprised to learn that robust estimates of the number of tropical tree species are lacking. We show that there are at least 40,000, but possibly more than 53,000, tree species in the tropics, in contrast to only 124 across temperate Europe. Almost all tropical tree species are restricted to their respective continents, and the Indo-Pacific region appears to be as species-rich as tropical America, with each of these two regions being almost five times as rich in tree species as African tropical forests. Our study shows that most tree species are extremely rare, meaning that they may be under serious risk of extinction at current deforestation rates. The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher’s alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between ∼40,000 and ∼53,000, i.e., at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of ∼19,000–25,000 tree species. Continental Africa is relatively depauperate with a minimum of ∼4,500–6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa.

Tree species stratification in relation to allometry and demography in a warm-temperate rain forest

1 We studied stratification of tree species by analysing the allometry and demography of 14 abundant species in a warm-temperate rain forest on Yakushima Island, southern Japan. 2 We regressed relationships between trunk diameter at breast height (d.b.h.) and top height of trees using an expanded allometry which incorporates an upper limit to height. As trees grew up towards the canopy, individuals with the same d.b.h. were taller if they came from species with larger maximum height. This appeared to reflect a trade-off between height growth and lateral growth of crowns. Two canopy dominants continued to gain in d.b.h. after they had reached the canopy, while their height remained almost constant. 3 We examined size-specific patterns of relative growth rate of d.b.h. from a decade of field censuses. By fitting a Gompertz-type growth curve to size-specific growth of individual trees, we derived a projected potential maximum size for each species. The projected maximum size (both d.b.h. and height) was similar to the observed value for most species. Three canopy species exhibited larger projected maximum d.b.h. than observed values; the difference seemed to be due to mortality during the long period that each of these species resides in the canopy. 4 Understorey species recruited equally well as larger-growing species amongst the 14 studied species, although their tree mass increments per unit time were smaller. This suggests that understorey species have more effective recruiting capacity relative to vegetative growth.

The effects of a typhoon on Japanese warm temperate rainforests

A powerful typhoon (Typhoon No. 13) affected stands of primary warm temperate rainforest in Yakushima, southern Japan, in 1993. Censuses in three sites 1–5 months after the typhoon showed low levels of mortality resulting from the typhoon (0.4–3.0% of stems). Stems killed by the typhoon were generally larger than surviving stems. Among surviving stems there were generally low levels of damage (e.g. 0.5–1.3% of surviving stems lost crowns) and not all damage was widespread (e.g. defoliation was apparent only in one of three sites). The sizes of uprooted stems, stems that lost crowns and relatively undamaged stems were not different. Different species appeared to be damaged in different ways at different sites. Damage was most evident in higher altitude seaward sites but gap formation was more frequent in lower altitude sites near valley floors. After the typhoon the resultant gap area occupied 9.4% of one site and 8.6% of another, which is greater than that before the typhoon. Fresh sprouts were found on 17.35% of 2161 stems after the typhoon, including many apparently undamaged stems as well as those that were damaged. Species which sprouted most frequently were those that regenerate by seed least frequently in these forests; these species may maintain their relative abundance in part by sprouting. Most tree species in these forests may be relatively resistant to typhoons and there may be more opportunities for their regeneration following gap formation caused by the typhoon.

Soil Phosphorus Fractionation and Phosphorus-Use Efficiency of a Bornean Tropical Montane Rain Forest During Soil Aging With Podozolization

We compared phosphorus (P) dynamics and plant productivity in two montane tropical rain forests (Mount Kinabalu, Borneo) that derived from similar parent materials (largely sedimentary rocks) and had similar climates but differed in terms of soil age. The younger site originated from Quaternary colluvial deposits, whereas the older site had Tertiary-age material. The older site had a distinctive spodic horizon, reduced levels of labile inorganic soil P, higher concentrations of recalcitrant organic soil P, and lower rates of net soil N mineralization. P fertilization led to soil nitrogen (N) immobilization in the P-deficient soil, indicating that soil N mineralization was limited by P at the P-deficient older site. Mean foliar nutrient concentration (on both a weight and an area basis) was similar at the two sites for all elements except P, which was lower at the older site. Aboveground net primary production (ANPP) was lower at the older site than at the younger one; this difference could be explained by the reduced availability of P and N (as down-regulated by P) at the older site. The relatively ample allocation of P and N to leaves, despite the reduced availability at the P-deficient old site, was attributable to its high resorption efficiency. High resorption resulted in lower concentrations of elements in leaf litter—that is, less decomposable low-quality litter. On the other hand, the concentration of leaf litter lignin was considerably lower at the older site; this appeared to be a de facto adaptive mechanism to avoid retarding litter decomposition.

Effects of the 1997–98 El Niño drought on rain forests of Mount Kinabalu, Borneo

We examined the effects of the 1997-98 El Nino drought on nine rain forests of Mount Kinabalu, Borneo, at four altitudes (700, 1700, 2700 and 3100 m) on contrasting geological substrata (ultrabasic versus non-ultrabasic). Measurements of rainfall and atmospheric aridity indicated that the departure from normal conditions during the drought became greater with increasing altitude. During 1997-99 (drought period) compared to 1995-97 (pre-drought period), median growth rates of stem diameter of trees decreased for both smaller (4.8-10 cm) and larger (≥ 10 cm) diameter classes in the six upland forcsts (≥ 2700 m on ultrabasic substrata and ≥ 1700 m on non-ultrabasic substrata), but for neither diameter class in the other forests. The majority of species decreased or did not change growth rates during 1997-99, whereas some did increase. Tree mortality increased during 1997-99, at the larger diameter class in the two lowland forests (700 m) on both substrata, and at least at the smaller diameter class in the four upland forests (≥ 1 700 m) on non-ultrabasic substrata. In two of these upland forests, mortality was restricted to particular understorey species. Mortality did not significantly increase in the three upland forests (≥ 1700 m) on ultrabasic substrata; this suggests that the adaptation to nutrient-poor soils might have provided the resistance to drought.

Changes in biomass, productivity and decomposition along topographical gradients under different geological conditions in tropical lower montane forests on Mount Kinabalu, Borneo

We have examined how the structure and function of a forest ecosystem change with topography (lower-slope versus ridge) and how the changes are modified by nutrient availability depending on geological substrate (Quaternary and Tertiary sedimentary rocks and ultrabasic rock) in the tropical montane rain forests of Mt. Kinabalu (Borneo) where climate is humid and aseasonal. Reflecting the difference in site age and parent rock, the pool size of soluble-P and inorganic-N in topsoils decreased from Quaternary sedimentary >Tertiary sedimentary >ultrabasic rock on the lower-slope, and they decreased from the lower-slope to the ridge on all substrates. Forest structural attributes [stature, above-ground biomass, and leaf area index (LAI)] decreased in the order of Quaternary sedimentary >Tertiary sedimentary >ultrabasic rock in association with soil nutrients on the lower-slopes, and decreased upslope consistently on each of the three substrates. Functional attributes [above-ground net primary productivity (ANPP) and decomposition rate] demonstrated similar patterns to structure. ANPP significantly correlated with LAI among the six sites, while net assimilation rate (ANPP divided by LAI assuming an even productivity between above vs below-ground system) was nearly constant. Therefore, ANPP could be explained primarily by LAI. Topographical change in LAI could be explained by leaf mass per area (LMA) combined with stand-level leaf biomass. LMA increased upslope on all substrates in association with the decrease in individual leaf area. Stand-level leaf biomass decreased upslope on all substrates but the Tertiary sedimentary rock. Our study demonstrated that topography and geological substrates interactively affected forest structure and processes. The effect of topography on forest structure and processes was greater on nutrient-rich substrates than on poor substrates, and the effect of geological substrate was greater on lower-slopes than on ridges.

Soils on exposed Sunda Shelf shaped biogeographic patterns in the equatorial forests of Southeast Asia

The marked biogeographic difference between western (Malay Peninsula and Sumatra) and eastern (Borneo) Sundaland is surprising given the long time that these areas have formed a single landmass. A dispersal barrier in the form of a dry savanna corridor during glacial maxima has been proposed to explain this disparity. However, the short duration of these dry savanna conditions make it an unlikely sole cause for the biogeographic pattern. An additional explanation might be related to the coarse sandy soils of central Sundaland. To test these two nonexclusive hypotheses, we performed a floristic cluster analysis based on 111 tree inventories from Peninsular Malaysia, Sumatra, and Borneo. We then identified the indicator genera for clusters that crossed the central Sundaland biogeographic boundary and those that did not cross and tested whether drought and coarse-soil tolerance of the indicator genera differed between them. We found 11 terminal floristic clusters, 10 occurring in Borneo, 5 in Sumatra, and 3 in Peninsular Malaysia. Indicator taxa of clusters that occurred across Sundaland had significantly higher coarse-soil tolerance than did those from clusters that occurred east or west of central Sundaland. For drought tolerance, no such pattern was detected. These results strongly suggest that exposed sandy sea-bed soils acted as a dispersal barrier in central Sundaland. However, we could not confirm the presence of a savanna corridor. This finding makes it clear that proposed biogeographic explanations for plant and animal distributions within Sundaland, including possible migration routes for early humans, need to be reevaluated.

Pattern of changes in species diversity, structure and dynamics of forest ecosystems along latitudinal gradients in East Asia

We examined effects of seasonality of climate and dominant life form (evergreen/deciduous, broad-leaf/coniferous) together with energy condition on species diversity, forest structure, forest dynamics, and productivity of forest ecosystems by comparing the patterns of changes in these ecosystem attributes along altitudinal gradients in tropical regions without seasonality and along a latitudinal gradient from tropical to temperate regions in humid East Asia. We used warmth index (temperature sum during growing season, WI) as an index of energy condition common to both altitudinal and latitudinal gradients. There were apparent differences in patterns of changes in the ecosystem attributes in relation to WI among four forest formations that were classified according to dominant life form and climatic zone (tropical/temperate). Many of the ecosystem attributes—Fisher’s alpha of species-diversity indices, maximum tree height and stem density, productivity [increment rate of aboveground biomass (AGB)], and population and biomass turnover rates—changed sharply with WI in tropical and temperate evergreen broad-leaved forests, but did not change linearly or changed only loosely with WI in temperate deciduous broad-leaved and evergreen coniferous forests. Values of these ecosystem attributes in temperate deciduous broad-leaved and evergreen coniferous forests were higher (stem density was lower) than those in tropical and temperate evergreen broad-leaved forests under colder conditions (WI below 100°C). Present results indicate that seasonality of climate and resultant change in dominant life form work to buffer the effects of energy reduction on ecosystem attributes along latitudinal gradients.

Effects of selective logging on tree species diversity and composition of Bornean tropical rain forests at different spatial scales

Reduced-impact logging (RIL) is known to be beneficial in biodiversity conservation, but its effects on tree diversity remain unknown. Pattern of tree diversity following disturbance usually varies with spatial scale of sampling (i.e., plot size). We examined the impacts of RIL on species richness and community composition of tree species at different spatial scales, and the scale (plot size) dependency of the two metrics; species richness versus community similarity. One 2-ha and three to four 0.2-ha plots were established in each of primary, RIL, and conventionally logged (CL) forest in Sabah, Malaysia. Species richness (the number of species per unit number of stems) was higher in the RIL than in the CL forest at both scales. The relationship between species richness and logging intensity varied with plot size. Species richness was greater in the RIL than in the primary forest at the 2-ha scale, while it was similar between the two forests at 0.2-ha scale. Similarly, species richness in the CL forest demonstrated a greater value at the 2-ha scale than at the 0.2-ha scale. Greater species richness in the two logged forests at the 2-ha scale is attributable to a greater probability of encountering the species-rich, small patches that are distributed heterogeneously. Community composition of the RIL forest more resembled that of the primary forest than that of the CL forest, regardless of plot size. Accordingly, species richness is a scale-dependent metric, while community similarity is a more robust metric to indicate the response of tree assemblage to anthropogenic disturbance.