Masaaki Takyu

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.

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.

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.

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.

Shoot growth and tree architecture of saplings of the major canopy dominants in a warm‐temperate rainforest

Sapling density, shoot growth, and sapling architecture were studied in five major canopy dominants both under closed canopy and gaps in a warm-temperate rainforest. The five species showed wide variations in distribution, shoot growth, and sapling architecture. Distylium racemosum and Quercus acuta had significantly higher sapling densities under closed canopy than in gaps. Castanopsis sieboldii and Machilus thunbergii had significantly higher sapling densities in gaps than under closed canopy. Quercus salicina showed no significant difference in sapling density between the two habitats. Under closed canopy, C. sieboldii and M. thunbergii had wider crowns than the other species. Distylium racemosum had the greatest number of terminal shoots among the species. Quercus acuta had a branchless small crown. Quercus salicina showed intermediate values in crown width, depth and the number of terminal shoots among the species. Distylium racemosum showed the greatest height-growth rate among the species under the closed canopy, but was the slowest in gaps. Castanopsis sieboldii and M. thunbergii showed the greatest height-growth rates among the species in gaps. Quercus salicina showed the slowest height-growth rates both under closed canopy and gaps. All of the five species showed low mortality under closed canopy. For the major canopy dominants: (i) sapling architecture may not be an important factor in determining mortality but it may be important for height-growth rate; and (ii) sprouting helps saplings to survive until gap formation and facilitates rapid growth in the gaps.

Community dynamics over 14 years along gradients of geological substrate and topography in tropical montane forests on Mount Kinabalu, Borneo

To understand the variation in community dynamics of tropical montane forests along gradients of soil fertility, death, recruitment and growth of trees ( 5 cm diameter) were monitored over 14 y (1997–2011) in nine plots placed in a matrix of three geological substrate types (Quaternary sediments, Tertiary sedimentary rocks and ultrabasic rocks) and three topographical units (ridge, middle and lower slopes) on Mount Kinabalu, Borneo. The plot area was 0.05 ha for ridge, 0.1 ha for middle slope and 0.2 ha (on ultrabasic rocks) and 1 ha (on the other substrates) for lower slope. Recruitment rates did not show a consistent pattern across geological substrates or topographies. Mortality rates were relatively high in almost all plots during the 1997–1999 period, including the El Niño drought, and in three plots on ultrabasic rocks during 2001–2005. Binomial logistic regression analyses showed that mortality during 1997–1999 increased with soil fertility (soluble phosphorus). Background mortality, excluding these periods, did not differ across geological substrates or topographies. The average growth rate during 1997–2011 was higher on more fertile soils and positively correlated with mortality during 1997–1999. We suggest that a high mortality rate during the drought period was related to high species diversity on more fertile soils, whereas a lower growth rate was related to stunted structures on poorer soils.

Structure, floristics and diversity of tropical montane rain forests over ultramafic soils on Mount Kinabalu (Borneo) compared with those on non-ultramafic soils

We describe here the structure, floristics and diversity of tropical montane rain forests over ultramafic soils on Mount Kinabalu, Borneo, and compared them with those on non-ultramafic soils. We used 14 sample plots from 1580 to 3080 m elevation, six on ultramafic soils and eight on non-ultramafic soils, and identified all trees ≥4.8 cm diameter. The plot area ranged from 0.1 to 1 ha, the majority (nine plots) being 0.25 ha. Forests on ultramafic soils showed more stunted structure, especially at higher altitudes, than those on non-ultramafic soils and on ridges than on slopes. Species of Coniferae (Araucariaceae and Podocarpaceae) and Myrtaceae strongly dominated on ultramafic soils occupying 61–96% of basal area in each plot, compared with 22–63% on non-ultramafic soils. Among 287 species found in the 14 plots, only nine species (including four species endemic to Mount Kinabalu) were strictly restricted to ultramafic soils. Nonmetric multidimensional scaling demonstrated that elevational change in species composition was accelerated on ultramafic soils and on ridges. Tree species diversity was generally lower on ultramafic soils than on non-ultramafic soils at the comparative altitudes. Multiple regression analysis suggested that soil nutrients (phosphorus and nitrogen) could be the cause of vegetation differentiation between ultramafic and non-ultramafic soils, although the data on soil metals are lacking. Comparison of our results with those from other mountains with ultramafic soils in South-east Asia demonstrated the uniqueness of the montane rain forests over ultramafic soils on Mount Kinabalu.

Management Effects on Tree Species Diversity and Dipterocarp Regeneration

Unregulated selective logging has been common throughout the Southeast Asian tropical rain forests. Such logging usually damages more than 50 % of the original forest biomass and causes soil disturbances by the use of heavy machinery (Cannon et al. 1994; Pinard and Putz 1996; Bertault and Sist 1997; Sist et al. 1998). To mitigate the deleterious logging impacts, reduced-impact logging has been applied in some tropical production forests (Kleine and Heuveldop 1993; Lagan et al. 2007; Putz et al. 2008a). Reduced-impact logging is a modification of selective logging that is intended to minimize collateral damage to the residual stands. Such methods include pre-harvest inventory, mapping of all canopy trees, directional felling, liana cutting, and planning of skid trails, log decks, and roads (see Chap. 1). In comparison with unregulated conventional logging, reduced-impact logging is beneficial in maintaining not only future crop trees (Rockwell et al. 2007; Pena-Claros et al. 2008) and forest biomass (Johns et al. 1996; Pinard and Putz 1996; Bertault and Sist 1997; Sist et al. 1998; Putz et al. 2008b; Imai et al. 2009), but also biological diversity, such as dung beetles (Davis 2000), flying insects (Akutsu et al. 2007), soil fauna (see Chap. 4 by Hasegawa et al., this volume), and forest-dwelling vertebrates (see Chap. 5 by Samejima et al., this volume). Many other taxa, such as ants, arachnids, bats, birds, fishes, and animals, also are not adversely affected by reduced-impact logging (Azevedo-Ramos et al. 2006; Wunderle et al. 2006; Castro-Arellano et al. 2007; Felton et al. 2008; Presley et al. 2008; Dias et al. 2010; Bicknell and Peres 2010). However, the effects of reduced-impact logging on tree species diversity of tropical rain forests remain largely unknown, despite that the diversity of trees is fundamental to the structure and functions of the forests. The diversity of trees may also determine the diversity of other taxonomic groups because trees provide resources and habitat structures for dependent species. The aim of this study is to examine the effects of reduced-impact logging versus conventional logging on tree species diversity and composition in the lowland tropical rain forests in Deramakot and Tangkulap.

Seasonal changes in pollen limitation and femaleness along the snowmelt gradient in a distylous alpine herb, Primula modesta

Abstract Flowering phenology of alpine plants is strongly determined by the timing of snowmelt, and the conditions of pollination of widely distributed plants vary greatly during their flowering season. We examined the reproductive success of the distylous alpine herb, Primula modesta, along the snowmelt gradient under natural conditions, and compared it with the result of artificial pollination experiments. In addition, the compositions and visit frequencies of pollinators to the flower of P. modesta were examined during the flowering period. The pin and thrum plants of P. modesta growing at the same site have an equal ability to produce seeds if a sufficient amount of legitimate pollen grains are deposited on the stigma surface. However, under natural conditions, their seed‐set success was often (even if not always) restricted by pollen limitation, and the functional gender of the pin and thrum plants biased to the female and male, respectively, associated with their growing sites. These variations were not ascribed to resource limitation nor biased morph ratio but to the seasonal changes in pollination situations, a replacement of pollinator types from long‐ to short‐tongued pollinators resulted in unidirectional pollen transfer from long stamens (thrum plants) to long styles (pin plants). The functional gender specialization may enhance the evolution of dioecy from heterostyly, but the severe pollen limitation may cause the breakdown of heterostyly into homostyly. To consider the evolutionary pathway of heterostylous plants, an accumulation of the empirical data is required demonstrating how phenological synchrony between plants and pollinators is decided and to what degree this relationship is stable over years, along with estimates of selection and gene flow in individual plants.