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.