Aim Large trees (d.b.h.≥70 cm) store large amounts of biomass. Several studies suggest that large trees may be vulnerable to changing climate, potentially leading to declining forest biomass storage. Here we determine the importance of large trees for tropical forest biomass storage and explore which intrinsic (species trait) and extrinsic (environment) variables are associated with the density of large trees and forest biomass at continental and pan-tropical scales. Location Pan-tropical. Methods Aboveground biomass (AGB) was calculated for 120 intact lowland moist forest locations. Linear regression was used to calculate variation in AGB explained by the density of large trees. Akaike information criterion weights (AICc-wi) were used to calculate averaged correlation coefficients for all possible multiple regression models between AGB/density of large trees and environmental and species trait variables correcting for spatial autocorrelation. Results Density of large trees explained c. 70% of the variation in pan-tropical AGB and was also responsible for significantly lower AGB in Neotropical [287.8 (mean)±105.0 (SD) Mg ha -1 versus Palaeotropical forests (Africa 418.3±91.8 Mg ha-1; Asia 393.3±109.3 Mg ha-1). Pan-tropical variation in density of large trees and AGB was associated with soil coarseness (negative), soil fertility (positive), community wood density (positive) and dominance of wind dispersed species (positive), temperature in the coldest month (negative), temperature in the warmest month (negative) and rainfall in the wettest month (positive), but results were not always consistent among continents. Main conclusions Density of large trees and AGB were significantly associated with climatic variables, indicating that climate change will affect tropical forest biomass storage. Species trait composition will interact with these future biomass changes as they are also affected by a warmer climate. Given the importance of large trees for variation in AGB across the tropics, and their sensitivity to climate change, we emphasize the need for in-depth analyses of the community dynamics of large trees. (Résumé d'auteur

Aim Tropical forests have been recognized as important global carbon sinks and sources. However, many uncertainties about the spatial distribution of live tree above-ground biomass (AGB) remain, mostly due to limited availability of AGB field data. Recent studies in the Amazon have already shown the importance of large sample size for accurate AGB gradient analysis. Here we use a large stem density, basal area, community wood density and AGB dataset to study and explain their spatial patterns in an Asian tropical forest.Location Borneo, Southeast Asia. MethodsWe combined stem density, basal area, community wood density and AGB data from 83 locations in Borneo with an environmental database containing elevation, climate and soil variables. The Akaike information criterion was used to select models and environmental variables that best explained the observed values of stem density, basal area, community wood density and AGB. These models were used to extrapolate these parameters across Borneo. ResultsWe found that wood density, stem density, basal area and AGB respond significantly, but differentially, to the environment. AGB was only correlated with basal area, but not with stem density and community wood specific gravity. Main conclusionsUnlike results from Amazonian forests, soil fertility was an important positive correlate for AGB in Borneo while community wood density, which is a main driver of AGB in the Neotropics, did not correlate with AGB in Borneo. Also, Borneo's average AGB of 457.1 Mg ha -1 was c. 60% higher than the Amazonian average of 288.6 Mg ha -1 . We find evidence that this difference might be partly explained by the high density of large wind-dispersed Dipterocarpaceae in Borneo, which need to be tall and emergent to disperse their seeds. Our results emphasize the importance of Bornean forests as carbon sinks and sources due to their high carbon storage capacity.

Summary1 Maximum attainable height varies greatly between tree species in tropical rain forests and covaries with demographic and allometric traits. We examined these relationships in 27 abundant tree species in a mixed dipterocarp forest. These species were monitored over 3 years in two 1-ha plots in western Borneo. A 95-percentile upper height limit was used to represent maximum height, to avoid sample size differences among populations. 2 Average growth rate in trunk diameter was regressed against trunk diameter using a maximum likelihood model and assuming that growth rates were exponentially distributed around the average. Estimated average growth rate at small trunk diameters (up to 11 cm) was independent of maximum height among the 27 species, while the degree of growth reduction at larger diameters was larger for species with smaller maximum height. 3 The recruitment rate efficiency of saplings was negatively correlated with maximum height, regardless of the measure used to assess species abundance. In particular, sapling recruitment per unit basal area declined greatly with increasing maximum height, consistent with model predictions of the traits required for the stable coexistence of species at different heights within the canopy. 4 Allometric analyses showed that understorey species had shorter heights at the same trunk diameter, and deeper crowns at the same tree height, than canopy species. Therefore, understorey species showed adaptive morphology to deep shade. 5 The regressed size-dependent pattern of average growth rate and an assumption that the population was in a steady state readily explained the observed trunk diameter distributions for 21 species among 27 examined. These species, for which the projected size distribution hardly changed when the natural increase or decrease of the population was set at γ = ± 0.005 year − 1 , had mortality rates of more than four times the value of γ .

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.

Wood properties were measured for trees in lowland dipterocarp forests in West Kalimantan. In 1993 and 1994, 353 samples of 286 species were collected from trunk base of trees of approximately 5 cm in diameter, and the specific gravities (SG: oven dry weight/fresh volume) and water contents of wood including bark were measured. The SG of each species ranged from 0.21 to 0.84, and the mean ± SD was 0.53 ± 0.13. The wide range of SG suggests that the forest had a high diversity in wood properties. The most dominant and diversified genus in this area was Shorea, and the SG of 15 species varied from 0.21 to 0.71. The range covered SG of pioneer (six Macaranga, 0.29–0.43) and small trees in primary forests (nine Eugenia and 10 Xanthophyllum, 0.55–0.77). The SG average for tree species of secondary forests of 2–6 years old was 0.31. It was significantly smaller than that of primary forests (0.58). In a primary dipterocarp forest plot, light‐wood species grew faster in diameter than heavy‐wood species. Water content ranged from 0.26 to 0.76. Heavy wood had low water content. Among light‐wood species, some (Shorea, Artocarpus) had low water contents and others (Ficus) had high water contents. Some riverine trees also had high water contents. These wood properties appear strongly related to the life history of trees and successional stage.

Aim  Identify environmental correlates for tropical tree diversity and composition. Location  Borneo, Southeast Asia. Methods  A GIS‐environmental database with 5 arc minute (c. 10 × 10 km) resolution was combined with tree inventory data. Tree diversity, phylogenetic diversity (PD) and the two main compositional gradients were determined for 46 tree inventories. Akaike's information criterion and a data jackknifing procedure were used to select 50 explanatory models for diversity and composition gradients. The average of these models was used as our final diversity and compositional model. We applied Moran's I to detect spatial autocorrelation of residuals. Results  Tree diversity, PD and the two main compositional gradients in Borneo were all significantly correlated with the environment. Tree diversity correlated negatively with elevation, soil depth, soil coarseness (texture) and organic carbon content, whereas it correlated positively with soil C:N ratio, soil pH, moisture storage capacity and annual rainfall. Tree PD was correlated positively with elevation and temperature seasonality and was largely determined by gymnosperms. However, angiosperm PD also correlated positive with elevation. Compositional patterns were strongly correlated with elevation but soil texture, cation‐exchange‐capacity, C:N ratio, C and N content and drainage were also important next to rainfall seasonality and El Niño Southern Oscillation drought impact. Main conclusions  Although elevation is the most important correlate for diversity and compositional gradients in Borneo, significant additional variability is explained by soil characteristics (texture, carbon content, pH, depth, drainage and nutrient status) and climate (annual rainfall, rainfall seasonality and droughts). The identified environmental correlates for diversity and composition gradients correspond to those found in other tropical regions of the world. Differences between the regions are mainly formed by differences in the relative importance of the environmental variables in explaining diversity and compositional gradients.

SignificanceIdentifying and explaining regional differences in tropical forest dynamics, structure, diversity, and composition are critical for anticipating region-specific responses to global environmental change. Floristic classifications are of fundamental importance for these efforts. Here we provide a global tropical forest classification that is explicitly based on community evolutionary similarity, resulting in identification of five major tropical forest regions and their relationships: (i) Indo-Pacific, (ii) Subtropical, (iii) African, (iv) American, and (v) Dry forests. African and American forests are grouped, reflecting their former western Gondwanan connection, while Indo-Pacific forests range from eastern Africa and Madagascar to Australia and the Pacific. The connection between northern-hemisphere Asian and American forests is confirmed, while Dry forests are identified as a single tropical biome.

Small-scale spatial association of the distribution for 55 abundant tree species with two environmental factors (humus depth and surface microtopography) was examined in two 1-ha plots of a heath (kerangas) forest in Central Kalimantan, Indonesia. More than 80% of the 55 species showed a significant habitat preference in humus depth and/or relative elevation in at least one plot. In particular, ten species occurring in both plots showed a consistent significant preference for humus depth or relative elevation in the two plots. Using randomization tests, however, only five species significantly associated with humus depth and no species with relative elevation. These results suggest that edaphic and topographic factors, especially humus depth, contribute to determining local spatial distribution and floristic composition of abundant tree species in the forest.