Tropical tree height-diameter (H:D) relationships may vary by forest type and region making large-scale estimates of above-ground biomass subject to bias if they ignore these differences in stem allometry. We have therefore developed a new global tropical forest database consisting of 39 955 concurrent H and D measurements encompassing 283 sites in 22 tropical countries. Utilising this database, our objectives were: 1. to determine if H:D relationships differ by geographic region and forest type (wet to dry forests, including zones of tension where forest and savanna overlap). 2. to ascertain if the H:D relationship is modulated by climate and/or forest structural characteristics (e.g. standlevel basal area, A). 3. to develop H:D allometric equations and evaluate biases to reduce error in future local-to-global estimates of tropical forest biomass. Annual precipitation coefficient of variation (PV), dry season length (SD), and mean annual air temperature (TA) emerged as key drivers of variation in H:D relationships at the pantropical and region scales. Vegetation structure also played a role with trees in forests of a high A being, on average, taller at any given D. After the effects of environment and forest structure are taken into account, two main regional groups can be identified. Forests in Asia, Africa and the Guyana Shield all have, on average, similar H:D relationships, but with trees in the forests of much of the Amazon Basin and tropical Australia typically being shorter at any given D than their counterparts elsewhere. The region-environment-structure model with the lowest Akaike's information criterion and lowest deviation estimated stand-level H across all plots to within a median -2.7 to 0.9% of the true value. Some of the plot-to-plot variability in H:D relationships not accounted for by this model could be attributed to variations in soil physical conditions. Other things being equal, trees tend to be more slender in the absence of soil physical constraints, especially at smaller D. Pantropical and continental-level models provided less robust estimates of H, especially when the roles of climate and stand structure in modulating H:D allometry were not simultaneously taken into account.Additional co-authors: T. F. Domingues, M. Drescher, P. M. Fearnside, M. B. Franca, N. M. Fyllas, G. Lopez-Gonzalez, A. Hladik, N. Higuchi, M. O. Hunter, Y. Iida, K. A. Salim, A. R. Kassim, M. Keller, J. Kemp, D. A. King, J. C. Lovett, B. S. Marimon, B. H. Marimon-Junior, E. Lenza, A. R. Marshall, D. J. Metcalfe, E. T. A. Mitchard, E. F. Moran, B.W. Nelson, R. Nilus, E. M. Nogueira, M. Palace, S. Patino, K. S.-H. Peh, M. T. Raventos, J. M. Reitsma, G. Saiz, F. Schrodt, B. Sonke, H. E. Taedoumg, S. Tan, H. Woll, and J. Lloy

Aboveground tropical tree biomass and carbon storage estimates commonly ignore tree height (<i>H</i>). We estimate the effect of incorporating <i>H</i> on tropics-wide forest biomass estimates in 327 plots across four continents using 42 656 <i>H</i> and diameter measurements and harvested trees from 20 sites to answer the following questions: <br><br> 1. What is the best <i>H</i>-model form and geographic unit to include in biomass models to minimise site-level uncertainty in estimates of destructive biomass? <br><br> 2. To what extent does including <i>H</i> estimates derived in (1) reduce uncertainty in biomass estimates across all 327 plots? <br><br> 3. What effect does accounting for <i>H</i> have on plot- and continental-scale forest biomass estimates? <br><br> The mean relative error in biomass estimates of destructively harvested trees when including <i>H</i> (mean 0.06), was half that when excluding <i>H</i> (mean 0.13). Power- and Weibull-<i>H</i> models provided the greatest reduction in uncertainty, with regional Weibull-<i>H</i> models preferred because they reduce uncertainty in smaller-diameter classes (&le;40 cm <i>D</i>) that store about one-third of biomass per hectare in most forests. Propagating the relationships from destructively harvested tree biomass to each of the 327 plots from across the tropics shows that including <i>H</i> reduces errors from 41.8 Mg ha⁻¹ (range 6.6 to 112.4) to 8.0 Mg ha⁻¹ (−2.5 to 23.0). For all plots, aboveground live biomass was −52.2 Mg ha⁻¹ (−82.0 to −20.3 bootstrapped 95% CI), or 13%, lower when including <i>H</i> estimates, with the greatest relative reductions in estimated biomass in forests of the Brazilian Shield, east Africa, and Australia, and relatively little change in the Guiana Shield, central Africa and southeast Asia. Appreciably different stand structure was observed among regions across the tropical continents, with some storing significantly more biomass in small diameter stems, which affects selection of the best height models to reduce uncertainty and biomass reductions due to <i>H</i>. After accounting for variation in <i>H</i>, total biomass per hectare is greatest in Australia, the Guiana Shield, Asia, central and east Africa, and lowest in east-central Amazonia, W. Africa, W. Amazonia, and the Brazilian Shield (descending order). Thus, if tropical forests span 1668 million km² and store 285 Pg C (estimate including <i>H</i>), then applying our regional relationships implies that carbon storage is overestimated by 35 Pg C (31–39 bootstrapped 95% CI) if <i>H</i> is ignored, assuming that the sampled plots are an unbiased statistical representation of all tropical forest in terms of biomass and height ...

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

HighlightsMicroclimate was monitored in primary forest, logged forest and oil palm plantation.There were strong relationships between leaf area index and diurnal climate.Logged forest is up to 2.5 °C warmer on average than primary forest.Oil palm plantations are up to 6.5 °C warmer on average than primary forest.Forest disturbance led to desiccation of the air near the forest floor.

Tropical forests are global centres of biodiversity and carbon storage. Many tropical countries aspire to protect forest to fulfil biodiversity and climate mitigation policy targets, but the conservation strategies needed to achieve these two functions depend critically on the tropical forest tree diversity-carbon storage relationship. Assessing this relationship is challenging due to the scarcity of inventories where carbon stocks in aboveground biomass and species identifications have been simultaneously and robustly quantified. Here, we compile a unique pan-tropical dataset of 360 plots located in structurally intact old-growth closed-canopy forest, surveyed using standardised methods, allowing a multi-scale evaluation of diversity-carbon relationships in tropical forests. Diversity-carbon relationships among all plots at 1 ha scale across the tropics are absent, and within continents are either weak (Asia) or absent (Amazonia, Africa). A weak positive relationship is detectable within 1 ha plots, indicating that diversity effects in tropical forests may be scale dependent. The absence of clear diversity-carbon relationships at scales relevant to conservation planning means that carbon-centred conservation strategies will inevitably miss many high diversity ecosystems. As tropical forests can have any combination of tree diversity and carbon stocks both require explicit consideration when optimising policies to manage tropical carbon and biodiversity.

Aim To test the extent to which the vertical structure of tropical forests is determined by environment, forest structure or biogeographical history. Location Pan‐tropical. Methods Using height and diameter data from 20,497 trees in 112 non‐contiguous plots, asymptotic maximum height (H AM) and height–diameter relationships were computed with nonlinear mixed effects (NLME) models to: (1) test for environmental and structural causes of differences among plots, and (2) test if there were continental differences once environment and structure were accounted for; persistence of differences may imply the importance of biogeography for vertical forest structure. NLME analyses for floristic subsets of data (only/excluding Fabaceae and only/excluding Dipterocarpaceae individuals) were used to examine whether family‐level patterns revealed biogeographical explanations of cross‐continental differences. Results H AM and allometry were significantly different amongst continents. H AM was greatest in Asian forests (58.3 ± 7.5 m, 95% CI), followed by forests in Africa (45.1 ± 2.6 m), America (35.8 ± 6.0 m) and Australia (35.0 ± 7.4 m), and height–diameter relationships varied similarly; for a given diameter, stems were tallest in Asia, followed by Africa, America and Australia. Precipitation seasonality, basal area, stem density, solar radiation and wood density each explained some variation in allometry and H AM yet continental differences persisted even after these were accounted for. Analyses using floristic subsets showed that significant continental differences in H AM and allometry persisted in all cases. Main conclusions Tree allometry and maximum height are altered by environmental conditions, forest structure and wood density. Yet, even after accounting for these, tropical forest architecture varies significantly from continent to continent. The greater stature of tropical forests in Asia is not directly determined by the dominance of the family Dipterocarpaceae, as on average non‐dipterocarps are equally tall. We hypothesise that dominant large‐statured families create conditions in which only tall species can compete, thus perpetuating a forest dominated by tall individuals from diverse families.

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.

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.