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 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.

Remote sensing is revolutionizing the way we study forests, and recent technological advances mean we are now able -for the first time -to identify and measure the crown dimensions of individual trees from airborne imagery. Yet to make full use of these data for quantifying forest carbon stocks and dynamics, a new generation of allometric tools which have tree height and crown size at their centre are needed. Here, we compile a global database of 108753 trees for which stem diameter, height and crown diameter have all been measured, including 2395 trees harvested to measure aboveground biomass. Using this database, we develop general allometric models for estimating both the diameter and aboveground biomass of trees from attributes which can be remotely sensed -specifically height and crown diameter. We show that tree height and crown diameter jointly quantify the aboveground biomass of individual trees and find that a single equation predicts stem diameter from these two variables across the world's forests. These new Correspondence: Tommaso Jucker, tel. +44 1223 333911, fax: +44 1223 333953,

Tree architecture, growth, and mortality change with increasing tree size and associated light conditions. To date, few studies have quantified how size-dependent changes in growth and mortality rates co-vary with architectural traits, and how such size-dependent changes differ across species and possible light capture strategies. We applied a hierarchical Bayesian model to quantify size-dependent changes in demographic rates and correlated demographic rates and architectural traits for 145 co-occurring Malaysian rain-forest tree species covering a wide range of tree sizes. Demographic rates were estimated using relative growth rate in stem diameter (RGR) and mortality rate as a function of stem diameter. Architectural traits examined were adult stature measured as the 95-percentile of the maximum stem diameter (upper diameter), wood density, and three tree architectural variables: tree height, foliage height, and crown width. Correlations between demographic rates and architectural traits were examined for stem diameters ranging from 1 to 47 cm. As a result, RGR and mortality varied significantly with increasing stem diameter across species. At smaller stem diameters, RGR was higher for tall trees with wide crowns, large upper diameter, and low wood density. Increased mortality was associated with low wood density at small diameters, and associated with small upper diameter and wide crowns over a wide range of stem diameters. Positive correlations between RGR and mortality were found over the whole range of stem diameters, but they were significant only at small stem diameters. Associations between architectural traits and demographic rates were strongest at small stem diameters. In the dark understory of tropical rain forests, the limiting amount of light is likely to make the interspecific difference in the effects of functional traits on demography more clear. Demographic performance is therefore tightly linked with architectural traits such as adult stature, wood density, and capacity for horizontal crown expansion. The enhancement of a demographic trade-off due to interspecific variation in functional traits in the understory helps to explain species coexistence in diverse rain forests.

Summary1. An important goal in plant community ecology is to understand how species traits determine demographic performance. Several functional traits have been shown to correlate with growth and mortality rates in trees, but less is known about how the relationships between functional traits and demographic rates change with tree size. 2. We examined the associations of functional traits with growth and mortality across 43 tree species in the Fushan 25-ha subtropical rain forest plot in northern Taiwan. We estimated the 95th percentile maximum stem diameter, wood density and six leaf functional traits (leaf area, specific leaf area, thickness, succulence, and mass-based nitrogen and phosphorus contents) obtained from leaves on juvenile and adult individuals of each species. 3. To quantify size-dependent changes in growth and mortality, relative growth rate (RGR) and mortality were estimated as a function of stem diameter using hierarchical Bayesian models. These rate estimates were then correlated with functional traits at a range of stem diameter classes. 4. Relationships between functional traits and demographic rates varied with tree size. Maximum size was positively correlated with RGR across a wide range of tree sizes. Wood density was negatively correlated with RGR and mortality for small-sized trees. Leaf traits such as leaf area and specific leaf area at juvenile and adult stages were associated more strongly with demographic rates for corresponding sizes than from other sizes. 5. Synthesis. The observed size-dependent changes in the trait-demography relationships are possibly due to the effects of developmental and environmental changes with increasing tree size. The underlying effects of functional traits on demographic performance vary with tree size, and this should influence dynamics in a tree community.

Summary1. Tree architecture is thought to allow species to partition horizontal and vertical light gradients in the forest canopy. Tree architecture is closely related to light capture, carbon gain and the efficiency with which trees reach the canopy. Previous studies that investigated how light gradients drive differentiation in tree architecture have produced inconsistent results, partially because of the differences in which tree species and ontogenetic stages were studied. 2. We examined the relationship between stem diameter, tree height, foliage height, crown width and life-history strategy over a broad size range of 200 randomly selected, co-occurring tree species in a lowland rainforest in Peninsular Malaysia. We developed a hierarchical Bayesian model to account for both intra-and interspecific variation and describe the relationships among tree architectural variables. We analysed interspecific variation in tree architectural variables in relation to adult stature and light requirement for species regeneration as a function of tree size. 3. There was little interspecific variation in architectural variables, this is partly because of large intraspecific variation in response to canopy heterogeneity, but it also suggests architectural convergence within this community. However, interspecific analyses showed that, for large-statured species, small size classes had thinner stems with narrow and shallow crowns, whereas large-size classes had wider crowns. Light-demanding species (as indicated by high sapling mortality in shaded conditions) showed weak trends in tree architecture and were only characterized by wide crowns at intermediate sizes.4. In summary, tree architectural traits overlapped across the species community. This suggests that architectural convergence and equalizing effects occur in this diverse tropical forest and that community-wide allometric equations can be used to describe forest height and carbon storage. Light resource partitioning also occurs, indicating stabilizing effects. Interspecific architectural variation in relation to adult stature supports the theory of the trade-off between early reproduction and vegetative growth. In closed rainforests, adult stature imposes a stronger force on architectural differentiation of species than regeneration light requirements.

Tropical tree height-diameter (<i>H:D</i>) 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 <i>H</i> and <i>D</i> measurements encompassing 283 sites in 22 tropical countries. Utilising this database, our objectives were: <br><br> &nbsp; 1. to determine if <i>H:D</i> relationships differ by geographic region and forest type (wet to dry forests, including zones of tension where forest and savanna overlap).<br><br> &nbsp; 2. to ascertain if the <i>H:D</i> relationship is modulated by climate and/or forest structural characteristics (e.g. stand-level basal area, <i>A</i>).<br><br> &nbsp; 3. to develop <i>H:D</i> allometric equations and evaluate biases to reduce error in future local-to-global estimates of tropical forest biomass. <br><br> Annual precipitation coefficient of variation (<i>P</i>_V), dry season length (<i>S</i>_D), and mean annual air temperature (<i>T</i>_A) emerged as key drivers of variation in <i>H:D</i> relationships at the pantropical and region scales. Vegetation structure also played a role with trees in forests of a high <i>A</i> being, on average, taller at any given <i>D</i>. 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 <i>H:D</i> relationships, but with trees in the forests of much of the Amazon Basin and tropical Australia typically being shorter at any given <i>D</i> than their counterparts elsewhere. <br><br> The region-environment-structure model with the lowest Akaike's information criterion and lowest deviation estimated stand-level <i>H</i> across all plots to within a median &ndash;2.7 to 0.9% of the true value. Some of the plot-to-plot variability in <i>H:D</i> 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 <i>D</i>. Pantropical and continental-level models provided only poor estimates of <i>H</i>, especially when the roles of climate and stand structure in modulating <i>H:D</i> allometry were not simultaneously taken into account