Patterns of both above-and belowground biomass and production were evaluated using published information from 200 individual data-sets. Data sets were comprised of the following types of information: organic matter storage in living and dead biomass (e.g. surface organic horizons and soil organic matter accumulations), aboveand belowground net primary production (NPP) and biomass, litter transfers, climatic data (i.e. precipitation and temperature), and nutrient storage (N, P, Ca, K) in above-and belowground biomass, soil organic matter and litter transfers. Forests were grouped by climate, foliage life-span, species and soil order. Several climatic and nutrient variables were regressed against fine root biomass or net primary production to determine what variables were most useful in predicting their dynamics. There were no significant or consistent patterns for above-and belowground biomass accumulation or NPP change across the different climatic forest types and by soil order. Similarly, there were no consistent patterns of soil organic matter (SOM) accumulation by climatic forest type but SOM varied significantly by soil order-the chemistry of the soil was more important in determining the amount of organic matter accumulation than climate. Soil orders which were high in aluminum, iron, and clay (e.g. Ultisols, Oxisols) had high total living and dead organic matter accumulations -especially in the cold temperate zone and in the tropics. Climatic variables and nutrient storage pools (i.e. in the forest floor) successfully predicted fine root NPP but not fine root biomass which was better predicted by nutrients in litterfall. The importance of grouping information by species based on their adaptive strategies for water and nutrient-use is suggested by the data. Some species groups did not appear to be sensitive to large changes in either climatic or nutrient variables while for others these variables explained a large proportion of the variation in fine root biomass and/or NPP.

1 Horizontal and vertical heterogeneity of resource availability, coupled with the specialized use of resources by tree species, results in complex patterns of tree species distributions in tropical rain forests. We studied the horizontal and vertical distributions of 4014 individuals in 11 species of early successional Macaranga (Euphorbiaceae) in tropical rain forest in Sarawak, Malaysia. 2 The horizontal distribution of individual trees was assessed with respect to crown light levels, establishment microsites, and broader scale variation in soil textural properties. Vertical distribution was assessed using an allometric approach to estimate maximum tree height (Hmax) and the slope of the sapling height–diameter relationship. 3 Average light levels intercepted and the proportion of individuals in each of five crown illumination classes varied significantly among the 11 species. Species ranged from extremely high‐light demanding, to quite shade tolerant. Average light levels intercepted by trees generally increased through ontogeny, but the ranking of species did not change significantly. 4 Fewer individuals of the more shade‐tolerant species established on disturbed microsites, irrespective of light levels. Among the more high‐light demanding species, the proportion of trees on different types of disturbed sites varied. 5 Trees of seven species were significantly more common on clay‐rich soils, two preferred sand‐rich soils, and two were not strongly affected by soil texture. 6 Hmax ranged from 5.5 to 31.3 m and was negatively correlated with shade tolerance among species, although among the more high‐light demanding species there was a wide range of tree sizes. Among species, Hmax was negatively correlated with both the slope and y‐intercept of the sapling height–diameter relationship, indicating that small‐statured species (also more shade tolerant) had more slender saplings than larger statured species. 7 Heterogeneity of resource availability leads to differences in horizontal and vertical tree distribution, which are important for the coexistence of 11 Macaranga species.

Summary 1We conducted a field experiment to test whether aggregated spatial distributions were related to soil variation in locally sympatric tree species in the rain forests of Sarawak, Malaysia. Dryobalanops aromatica , Shorea laxa , and Swintonia schwenkii are naturally aggregated on low-fertility humult ultisols, Dryobalanops lanceolata and Hopea dryobalanoides on moderate-fertility udult ultisols and Shorea balanocarpoides is found on both soil types. 2 Seedlings of all six species were grown in a nested-factorial experiment for 20 months in humult and udult soils in gaps and in the understorey to test for soil-specific differences in performance. Phosphorus addition was used to test for effects due to P-limitation. 3 Four species showed significantly higher growth on their natural soils, but one humultsoil species ( D. aromatica ) and the broadly distributed species were not significantly affected by soil type. 4 One udult-soil species, D. lanceolata , had both lower relative growth rate and lower mycorrhizal colonization on humult soil. However, humult soils also had lower levels of Ca, Mg, K, N and probably water availability. 5 The overall ranking of growth rates among species was similar on the two soils. Growth rates were strongly positively correlated with leaf area ratio and specific leaf area among species in both soils. With the exception of D. aromatica , species of the higher-nutrient soils had higher growth rates on both soils. 6 Although P addition led to elevated soil-P concentrations, elevated root-and leaf-tissue P concentrations on both soils, there was no significant growth enhancement and therefore no evidence that P availability limits the growth or constrains the distribution of any of the six species in the field. Differences in soil water availability between soils may be more important. 7 Our results suggest that habitat-mediated differences in seedling performance strongly influence the spatial distributions of tropical trees and are therefore likely to play a key role in structuring tropical rain forest communities.

Discrete humus layers are common on podzols under temperate coniferous and tropical heath forests, and patchy layers also occur under some temperate broadleaved forests on non-podzolic soils. We used multiple data sets to test the reported association of humus with oligotrophic but non-podzolic soils under non-heath dipterocarp forest at Lambir, Sarawak. We examined the distribution, morphology and nutrient dynamics of necromass on soils derived from sandstone and shale. Concentrations of the main mineral nutrients were lower in fresh litter on the very oligotrophic sandstone soils than on shale. The rates of litterfall were similar, so that annual litterfall fluxes of all nutrients were lower on sandstone. The lower nutrient concentrations and fluxes in the litter on sandstone resulted in slower decomposition, longer residence times and larger standing crops of forest-floor necromass, with lower concentrations of nutrients. The necromass on sandstone sequestered significantly more N, K and Mg but less Ca and Mn than on shale, with no significant difference for P. The variations in necromass nutrient dynamics were associated with morphological differences. There were mats of densely rooted humus under the litter on sandstone, whereas litter lay directly over the mineral topsoil on shale. Spatial associations with soil nutrients were weak for necromass thickness, but clear for humus. The proportions of nutrients in the litterfall and necromass reflected the stoichiometric profiles of the soils. We attribute the differences in necromass nutrient dynamics and their association with soil reserve nutrients to lower rates of nutrient replenishment from the weathering of sandstone than from shale. Necromass characteristics are robust field indicators of multivariate edaphic differences in these and other tropical forests on Acrisols/Ultisols derived from Tertiary clastic sediments.

Relationships between spatial distributions and site conditions, namely topography and soil texture, were analyzed for two congeneric emergent trees, Dryobalanops aromatica and Dryobalanops lanceolata (Dipterocarpaceae), in a tropical rainforest in Sarawak, East Malaysia. A 52‐ha permanent plot was divided into 1300 quadrats measuring 20 m × 20 m; for each Dryobalanops species, the number and total basal area of trees ≥1 cm in d.b.h. were compared among groups of quadrats with different site conditions. Because spatial distributions of both Dryobalanops and site‐condition variables were aggregated, Monte‐Carlo permutation tests were applied to analyze the relationships. Both single and multifactor statistical tests showed that the density and basal area distributions of the two species were significantly non‐random in relation to soil texture and topographic variables. D. aromatica was significantly more abundant at higher elevations, in sandy soils, and on convex and steep slopes. In contrast, D. lanceolata preferred lower elevations and less sandy soils. In the study plot, there were very few sites (3 of 1150 quadrats tested) where the models of Hayashi's method predicted the co‐occurrence of the two species. These results suggest that between‐species differences in habitat preferences are so large that they alone explain the spatially segregated distributions of these two species within the 52‐ha study plot.

The approximately 300 (298, 95% CI: 152–581) elephants in the Lower Kinabatangan Managed Elephant Range in Sabah, Malaysian Borneo are a priority sub-population for Borneo's total elephant population (2,040, 95% CI: 1,184–3,652). Habitat loss and human-elephant conflict are recognized as the major threats to Bornean elephant survival. In the Kinabatangan region, human settlements and agricultural development for oil palm drive an intense fragmentation process. Electric fences guard against elephant crop raiding but also remove access to suitable habitat patches. We conducted expert opinion-based least-cost analyses, to model the quantity and configuration of available suitable elephant habitat in the Lower Kinabatangan, and called this the Elephant Habitat Linkage. At 184 km2, our estimate of available habitat is 54% smaller than the estimate used in the State's Elephant Action Plan for the Lower Kinabatangan Managed Elephant Range (400 km2). During high flood levels, available habitat is reduced to only 61 km2. As a consequence, short-term elephant densities are likely to surge during floods to 4.83 km−2 (95% CI: 2.46–9.41), among the highest estimated for forest-dwelling elephants in Asia or Africa. During severe floods, the configuration of remaining elephant habitat and the surge in elephant density may put two villages at elevated risk of human-elephant conflict. Lower Kinabatangan elephants are vulnerable to the natural disturbance regime of the river due to their limited dispersal options. Twenty bottlenecks less than one km wide throughout the Elephant Habitat Linkage, have the potential to further reduce access to suitable habitat. Rebuilding landscape connectivity to isolated habitat patches and to the North Kinabatangan Managed Elephant Range (less than 35 km inland) are conservation priorities that would increase the quantity of available habitat, and may work as a mechanism to allow population release, lower elephant density, reduce human-elephant conflict, and enable genetic mixing.

No abstract

The diversity and niche specificity of hemi‐epiphytic figs in a lowland dipterocarp forest in Sarawak were investigated in 1998. Twenty‐seven fig species (264 individuals, c. 120 ha) colonized a diversity of host taxa (35 families), but densities were very low and only 1.77% of trees> 30 cm d.b.h. were occupied. There were no significant associations with host taxa or host‐bark roughness but among 11 common species (≥9 individuals) the distributions of all other parameters (host‐d.b.h., height and position of colonization, crown illumination, soil‐texture and slope‐angle) were significantly different, and we identified five fig guilds. The guilds corresponded to canopy strata, and appeared to reflect   the establishment microsite requirements of different species. A fundamental trade‐off within the hemi‐epiphytic habit was revealed: Species colonizing larger hosts were rarer, because of lower host densities and more specific microsite requirements, but had better light environments and attained a larger maximum size. The single strangler species appeared to escape many of these constraints, and an important source of mortality caused by host‐toppling, indicating the advantages of this strategy. Thus, the hemi‐epiphytic figs in this community have come to fill a remarkable diversity of niches, despite low levels of competition, through the exigencies of a complex environment. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78, 439–455