Damage to living trees contributes to almost half of the biomass losses in tropical forests

Accurate estimates of forest biomass stocks and fluxes are needed to quantify global carbon budgets and assess the response of forests to climate change. However, most forest inventories consider tree mortality as the only aboveground biomass (AGB) loss without accounting for losses via damage to living trees: branchfall, trunk breakage, and wood decay. Here, we use ~151,000 annual records of tree survival and structural completeness to compare AGB loss via damage to living trees to total AGB loss (mortality + damage) in seven tropical forests widely distributed across environmental conditions. We find that 42% (3.62 Mg ha⁻¹ year⁻¹; 95% confidence interval [CI] 2.36–5.25) of total AGB loss (8.72 Mg ha⁻¹ year⁻¹; CI 5.57–12.86) is due to damage to living trees. Total AGB loss was highly variable among forests, but these differences were mainly caused by site variability in damage-related AGB losses rather than by mortality-related AGB losses. We show that conventional forest inventories overestimate stand-level AGB stocks by 4% (1%–17% range across forests) because assume structurally complete trees, underestimate total AGB loss by 29% (6%–57% range across forests) due to overlooked damage-related AGB losses, and overestimate AGB loss via mortality by 22% (7%–80% range across forests) because of the assumption that trees are undamaged before dying. Our results indicate that forest carbon fluxes are higher than previously thought. Damage on living trees is an underappreciated component of the forest carbon cycle that is likely to become even more important as the frequency and severity of forest disturbances increase.     

Variation in wood density determines spatial patterns inAmazonian forest biomass

Uncertainty in biomass estimates is one of the greatest limitations to models of carbon flux in tropical forests. Previous comparisons of field‐based estimates of the aboveground biomass (AGB) of trees greater than 10 cm diameter within Amazonia have been limited by the paucity of data for western Amazon forests, and the use of site‐specific methods to estimate biomass from inventory data. In addition, the role of regional variation in stand‐level wood specific gravity has not previously been considered. Using data from 56 mature forest plots across Amazonia, we consider the relative roles of species composition (wood specific gravity) and forest structure (basal area) in determining variation in AGB. Mean stand‐level wood specific gravity, on a per stem basis, is 15.8% higher in forests in central and eastern, compared with northwestern Amazonia. This pattern is due to the higher diversity and abundance of taxa with high specific gravity values in central and eastern Amazonia, and the greater diversity and abundance of taxa with low specific gravity values in western Amazonia. For two estimates of AGB derived using different allometric equations, basal area explains 51.7% and 63.4%, and stand‐level specific gravity 45.4% and 29.7%, of the total variation in AGB. The variation in specific gravity is important because it determines the regional scale, spatial pattern of AGB. When weighting by specific gravity is included, central and eastern Amazon forests have significantly higher AGB than stands in northwest or southwest Amazonia. The regional‐scale pattern of species composition therefore defines a broad gradient of AGB across Amazonia.     

Deadwood stocks increase with selective logging and large tree frequency in Gabon

Deadwood is a major component of aboveground biomass (AGB) in tropical forests and is important as habitat and for nutrient cycling and carbon storage. With deforestation and degradation taking place throughout the tropics, improved understanding of the magnitude and spatial variation in deadwood is vital for the development of regional and global carbon budgets. However, this potentially important carbon pool is poorly quantified in Afrotropical forests and the regional drivers of deadwood stocks are unknown. In the first large‐scale study of deadwood in Central Africa, we quantified stocks in 47 forest sites across Gabon and evaluated the effects of disturbance (logging), forest structure variables (live AGB, wood density, abundance of large trees), and abiotic variables (temperature, precipitation, seasonality). Average deadwood stocks (measured as necromass, the biomass of deadwood) were 65 Mg ha^?1 or 23% of live AGB. Deadwood stocks varied spatially with disturbance and forest structure, but not abiotic variables. Deadwood stocks increased significantly with logging (+38 Mg ha^?1) and the abundance of large trees (+2.4 Mg ha^?1 for every tree >60 cm dbh). Gabon holds 0.74 Pg C, or 21% of total aboveground carbon in deadwood, a threefold increase over previous estimates. Importantly, deadwood densities in Gabon are comparable to those in the Neotropics and respond similarly to logging, but represent a lower proportion of live AGB (median of 18% in Gabon compared to 26% in the Neotropics). In forest carbon accounting, necromass is often assumed to be a constant proportion (9%) of biomass, but in humid tropical forests this ratio varies from 2% in undisturbed forest to 300% in logged forest. Because logging significantly increases the deadwood carbon pool, estimates of tropical forest carbon should at a minimum use different ratios for logged (mean of 30%) and unlogged forests (mean of 18%).     

Rapid Simultaneous Estimation of Aboveground Biomass and Tree Diversity Across Neotropical Forests: A Comparison of Field Inventory Methods

A standardized rapid inventory method providing information on both tree species diversity and aboveground carbon stocks in tropical forests will be an important tool for evaluating efforts to conserve biodiversity and to estimate the carbon emissions that result from deforestation and degradation (REDD). Herein, we contrast five common plot methods differing in shape, size, and effort requirements to estimate tree diversity and aboveground tree biomass (AGB). We simulated the methods across six Neotropical forest sites that represent a broad gradient in forest structure, tree species richness, and floristic composition, and we assessed the relative performance of methods by evaluating the bias and precision of their estimates of AGB and tree diversity. For a given sample of forest area, a ‘several small’ (< 1 ha) sampling strategy led to a smaller coefficient of variation (CV) in the estimate of AGB than a ‘few large’ one. The effort (person‐days) required to achieve an accurate AGB estimate (< 10% CV), however, was greater for the smallest plots (0.1 ha) than for a compromise approach using 0.5 ha modified Gentry plots, which proved to be the most efficient method to estimate AGB across all forest types. Gentry plots were also the most efficient at providing accurate estimates of tree diversity (< 10% CV of Hill number). We recommend the use of the 0.5 ha modified Gentry plot method in future rapid inventories, and we discuss a set of criteria that should inform any choice of inventory method.     

Simulating Carbon Stocks and Fluxes of an African Tropical Montane Forest with an Individual-Based Forest Model

Tropical forests are carbon-dense and highly productive ecosystems. Consequently, they play an important role in the global carbon cycle. In the present study we used an individual-based forest model (FORMIND) to analyze the carbon balances of a tropical forest. The main processes of this model are tree growth, mortality, regeneration, and competition. Model parameters were calibrated using forest inventory data from a tropical forest at Mt. Kilimanjaro. The simulation results showed that the model successfully reproduces important characteristics of tropical forests (aboveground biomass, stem size distribution and leaf area index). The estimated aboveground biomass (385 t/ha) is comparable to biomass values in the Amazon and other tropical forests in Africa. The simulated forest reveals a gross primary production of 24 t_cha^-1yr^-1. Modeling above- and belowground carbon stocks, we analyzed the carbon balance of the investigated tropical forest. The simulated carbon balance of this old-growth forest is zero on average. This study provides an example of how forest models can be used in combination with forest inventory data to investigate forest structure and local carbon balances.     

Mass‐ratio and complementarity effects simultaneously drive aboveground biomass in temperate Quercus forests through stand structure

Forests play a key role in regulating the global carbon cycle, a substantial portion of which is stored in aboveground biomass (AGB). It is well understood that biodiversity can increase the biomass through complementarity and mass‐ratio effects, and the contribution of environmental factors and stand structure attributes to AGB was also observed. However, the relative influence of these factors in determining the AGB of Quercus forests remains poorly understood. Using a large dataset retrieved from 523 permanent forest inventory plots across Northeast China, we examined the effects of integrated multiple tree species diversity components (i.e., species richness, functional, and phylogenetic diversity), functional traits composition, environmental factors (climate and soil), stand age, and structure attributes (stand density, tree size diversity) on AGB based on structural equation models. We found that species richness and phylogenetic diversity both were not correlated with AGB. However, functional diversity positively affected AGB via an indirect effect in line with the complementarity effect. Moreover, the community‐weighted mean of specific leaf area and height increased AGB directly and indirectly, respectively; demonstrating the mass‐ratio effect. Furthermore, stand age, density, and tree size diversity were more important modulators of AGB than biodiversity. Our study highlights that biodiversity–AGB interaction is dependent on the regulation of stand structure that can be even more important for maintaining high biomass than biodiversity in temperate Quercus forests.     

Biomass dynamics in a logged forest: the role of wood density

Wood density (WD) is believed to be a key trait in driving growth strategies of tropical forest species, and as it entails the amount of mass per volume of wood, it also tends to correlate with forest carbon stocks. Yet there is relatively little information on how interspecific variation in WD correlates with biomass dynamics at the species and population level. We determined changes in biomass in permanent plots in a logged forest in Vietnam from 2004 to 2012, a period representing the last 8 years of a 30 years logging cycle. We measured diameter at breast height (DBH) and estimated aboveground biomass (AGB) growth, mortality, and net AGB increment (the difference between AGB gains and losses through growth and mortality) per species at the individual and population (i.e. corrected for species abundance) level, and correlated these with WD. At the population level, mean net AGB increment rates were 6.47 Mg ha^?1 year^?1 resulting from a mean AGB growth of 8.30 Mg ha^?1 year^?1, AGB recruitment of 0.67 Mg ha^?1 year^?1 and AGB losses through mortality of 2.50 Mg ha^?1 year^?1. Across species there was a negative relationship between WD and mortality rate, WD and DBH growth rate, and a positive relationship between WD and tree standing biomass. Standing biomass in turn was positively related to AGB growth, and net AGB increment both at the individual and population level. Our findings support the view that high wood density species contribute more to total biomass and indirectly to biomass increment than low wood density species in tropical forests. Maintaining high wood density species thus has potential to increase biomass recovery and carbon sequestration after logging.     

Drivers of aboveground wood production in a lowland tropical forest of West Africa: teasing apart the roles of tree density,tree diversity,soil phosphorus,and historical logging

Tropical forests currently play a key role in regulating the terrestrial carbon cycle and abating climate change by storing carbon in wood. However, there remains considerable uncertainty as to whether tropical forests will continue to act as carbon sinks in the face of increased pressure from expanding human activities. Consequently, understanding what drives productivity in tropical forests is critical. We used permanent forest plot data from the Gola Rainforest National Park (Sierra Leone) – one of the largest tracts of intact tropical moist forest in West Africa – to explore how (1) stand basal area and tree diversity, (2) past disturbance associated with past logging, and (3) underlying soil nutrient gradients interact to determine rates of aboveground wood production (AWP). We started by statistically modeling the diameter growth of individual trees and used these models to estimate AWP for 142 permanent forest plots. We then used structural equation modeling to explore the direct and indirect pathways which shape rates of AWP. Across the plot network, stand basal area emerged as the strongest determinant of AWP, with densely packed stands exhibiting the fastest rates of AWP. In addition to stand packing density, both tree diversity and soil phosphorus content were also positively related to productivity. By contrast, historical logging activities negatively impacted AWP through the removal of large trees, which contributed disproportionately to productivity. Understanding what determines variation in wood production across tropical forest landscapes requires accounting for multiple interacting drivers – with stand structure, tree diversity, and soil nutrients all playing a key role. Importantly, our results also indicate that logging activities can have a long‐lasting impact on a forest's ability to sequester and store carbon, emphasizing the importance of safeguarding old‐growth tropical forests.     

Effects of endogenous and exogenous processes on aboveground biomass stocks and dynamics in Andean forests

Tropical forests are paramount in regulating the global carbon cycle due to the storage of large amounts of carbon in their biomass. Using repeat censuses of permanent plots located at 15 sites in the Andes Mountains of northwest Colombia, we evaluate: (1) the relationship between aboveground biomass (AGB) stocks, AGB dynamics (mortality, productivity, and net change), and changes in temperature across a ca. 3000-m elevational gradient (≈?16.1 °C); (2) how AGB mortality and AGB productivity interact to determine net AGB change; and (3) the extent to which either fine-grain (0.04-ha) or coarse-grain (1-ha) processes determine the AGB dynamics of these forests. We did not find a significant relationship between elevation/temperature and biomass stocks. The net AGB sequestered each year by these forests (2.21?±?0.51 Mg ha^?1 year^?1), equivalent to approximately 1.09% of initial AGB, was primarily determined by tree growth. Both forest structural properties and global warming influenced AGB mortality and net change. AGB productivity increases with greater inequality of tree sizes, a pattern characteristic of forest patches recovering from disturbances. Overall, we find that global warming is triggering directional changes in species composition by thermophilization via increased tree mortality of species in the lower portions of their thermal ranges and that the inclusion of small-scale forest structural changes can effectively account for endogenous processes such as changes in forest structure. The inclusion of fine-grain processes in assessments of AGB dynamics could provide additional insights about the effects that ongoing climate change has on the functioning of tropical montane forests.     

Improved allometric models to estimate the aboveground biomass of tropical trees

Terrestrial carbon stock mapping is important for the successful implementation of climate change mitigation policies. Its accuracy depends on the availability of reliable allometric models to infer oven‐dry aboveground biomass of trees from census data. The degree of uncertainty associated with previously published pantropical aboveground biomass allometries is large. We analyzed a global database of directly harvested trees at 58 sites, spanning a wide range of climatic conditions and vegetation types (4004 trees ≥ 5 cm trunk diameter). When trunk diameter, total tree height, and wood specific gravity were included in the aboveground biomass model as covariates, a single model was found to hold across tropical vegetation types, with no detectable effect of region or environmental factors. The mean percent bias and variance of this model was only slightly higher than that of locally fitted models. Wood specific gravity was an important predictor of aboveground biomass, especially when including a much broader range of vegetation types than previous studies. The generic tree diameter–height relationship depended linearly on a bioclimatic stress variable E, which compounds indices of temperature variability, precipitation variability, and drought intensity. For cases in which total tree height is unavailable for aboveground biomass estimation, a pantropical model incorporating wood density, trunk diameter, and the variable E outperformed previously published models without height. However, to minimize bias, the development of locally derived diameter–height relationships is advised whenever possible. Both new allometric models should contribute to improve the accuracy of biomass assessment protocols in tropical vegetation types, and to advancing our understanding of architectural and evolutionary constraints on woody plant development.     

Optimal climate for large trees at high elevations drives patterns of biomass in remote forests of Papua New Guinea

Our ability to model global carbon fluxes depends on understanding how terrestrial carbon stocks respond to varying environmental conditions. Tropical forests contain the bulk of the biosphere's carbon. However, there is a lack of consensus as to how gradients in environmental conditions affect tropical forest carbon. Papua New Guinea (PNG) lies within one of the largest areas of contiguous tropical forest and is characterized by environmental gradients driven by altitude; yet, the region has been grossly understudied. Here, we present the first field assessment of aboveground biomass (AGB) across three main forest types of PNG using 193 plots stratified across 3,100‐m elevation gradient. Unexpectedly, AGB had no direct relationship to rainfall, temperature, soil, or topography. Instead, natural disturbances explained most variation in AGB. While large trees (diameter at breast height > 50 cm) drove altitudinal patterns of AGB, resulting in a major peak in AGB (2,200–3,100 m) and some of the most carbon‐rich forests at these altitudes anywhere. Large trees were correlated to a set of climatic variables following a hump‐shaped curve. The set of “optimal” climatic conditions found in montane cloud forests is similar to that of maritime temperate areas that harbor the largest trees in the world: high ratio of precipitation to evapotranspiration (2.8), moderate mean annual temperature (13.7°C), and low intra‐annual temperature range (7.5°C). At extreme altitudes (2,800–3,100 m), where tree diversity elsewhere is usually low and large trees are generally rare or absent, specimens from 18 families had girths >70 cm diameter and maximum heights 20–41 m. These findings indicate that simple AGB‐climate‐edaphic models may not be suitable for estimating carbon storage in forests where optimal climate niches exist. Our study, conducted in a very remote area, suggests that tropical montane forests may contain greater AGB than previously thought and the importance of securing their future under a changing climate is therefore enhanced.     

Variation in above‐ground forest biomass across broad climatic gradients

Aim An understanding of the relationship between forest biomass and climate is needed to predict the impacts of climate change on carbon stores. Biomass patterns have been characterized at geographically or climatically restricted scales, making it unclear if biomass is limited by climate in any general way at continental to global scales. Using a dataset spanning multiple climatic regions we evaluate the generality of published biomass–climate correlations. We also combine metabolic theory and hydraulic limits to plant growth to first derive and then test predictions for how forest biomass should vary with maximum individual tree biomass and the ecosystem water deficit. Location Temperate forests and dry, moist and wet tropical forests across North, Central and South America. Methods A forest biomass model was derived from allometric functions and power‐law size distributions. Biomass and climate were correlated using extensive forest plot (276 0.1‐ha plots), wood density and climate datasets. Climate variables included mean annual temperature, annual precipitation, their ratio, precipitation of the driest quarter, potential and actual evapotranspiration, and the ecosystem water deficit. The water deficit uniquely summarizes water balance by integrating water inputs from precipitation with water losses due to solar energy. Results Climate generally explained little variation in forest biomass, and mixed support was found for published biomass–climate relationships. Our theory indicated that maximum individual biomass governs forest biomass and is constrained by water deficit. Indeed, forest biomass was tightly coupled to maximum individual biomass and the upper bound of maximum individual biomass declined steeply with water deficit. Water deficit similarly constrained the upper bound of forest biomass, with most forests below the constraint. Main conclusions The results suggest that: (1) biomass–climate models developed at restricted geographic/climatic scales may not hold at broader scales; (2) maximum individual biomass is strongly related to forest biomass, suggesting that process‐based models should focus on maximum individual biomass; (3) the ecosystem water deficit constrains biomass, but realized biomass often falls below the constraint; such that (4) biomass is not strongly limited by climate in most forests so that forest biomass may not predictably respond to changes in mean climate.     

Forest aboveground biomass along an elevational transect in Sulawesi,Indonesia, and the role of Fagaceae in tropical montane rain forests

Aim This study investigates how estimated tree aboveground biomass (AGB) of tropical montane rain forests varies with elevation, and how this variation is related to elevational change in floristic composition, phylogenetic community structure and the biogeography of the dominant tree taxa. Location Lore Lindu National Park, Sulawesi, Indonesia. Methods Floristic inventories and stand structural analyses were conducted on 13 plots (each 0.24 ha) in four old‐growth forest stands at 1050, 1400, 1800 and 2400 m a.s.l. (submontane to upper montane elevations). Tree AGB estimates were based on d.b.h., height and wood specific gravity. Phylogenetic diversity and biogeographical patterns were analysed based on tree family composition weighted by AGB. Elevational trends in AGB were compared with other Southeast Asian and Neotropical transect studies (n = 7). Results AGB was invariant from sub‐ to mid‐montane elevation (309–301 Mg ha^?1) and increased slightly to 323 Mg ha^?1 at upper montane elevation. While tree and canopy height decreased, wood specific gravity increased. Magnoliids accounted for most of the AGB at submontane elevations, while eurosids I (including Fagaceae) contributed substantially to AGB at all elevations. Phylogenetic diversity was highest at upper montane elevations, with co‐dominance of tree ferns, Podocarpaceae, Trimeniaceae and asterids/euasterids II, and was lowest at lower/mid‐montane elevations, where Fagaceae contributed > 50% of AGB. Biogeographical patterns showed a progression from dominant tropical families at submontane to tropical Fagaceae (Castanopsis, Lithocarpus) at lower/mid‐montane, and to conifers and Australasian endemics at upper montane elevations. Cross‐continental comparisons revealed an elevational AGB decrease in transects with low/no presence of Fagaceae, but relatively high AGB in montane forests with moderate to high abundance of this family. Main conclusions AGB is determined by both changes in forest structure and shifts in species composition. In our study, these two factors traded off so that there was no net change in AGB, even though there were large changes in forest structure and composition along the elevational gradient. Southeast Asian montane rain forests dominated by Fagaceae constitute important carbon stocks. The importance of biogeography and species traits for biomass estimation should be considered by initiatives to reduce emissions from deforestation and forest degradation (REDD) and in taxon choice in reforestation for carbon offsetting.     

The regional variation of aboveground live biomass in old-growth Amazonian forests

The biomass of tropical forests plays an important role in the global carbon cycle, both as a dynamic reservoir of carbon, and as a source of carbon dioxide to the atmosphere in areas undergoing deforestation. However, the absolute magnitude and environmental determinants of tropical forest biomass are still poorly understood. Here, we present a new synthesis and interpolation of the basal area and aboveground live biomass of old‐growth lowland tropical forests across South America, based on data from 227 forest plots, many previously unpublished. Forest biomass was analyzed in terms of two uncorrelated factors: basal area and mean wood density. Basal area is strongly affected by local landscape factors, but is relatively invariant at regional scale in moist tropical forests, and declines significantly at the dry periphery of the forest zone. Mean wood density is inversely correlated with forest dynamics, being lower in the dynamic forests of western Amazonia and high in the slow‐growing forests of eastern Amazonia. The combination of these two factors results in biomass being highest in the moderately seasonal, slow growing forests of central Amazonia and the Guyanas (up to 350 Mg dry weight ha^?1) and declining to 200–250 Mg dry weight ha^?1 at the western, southern and eastern margins. Overall, we estimate the total aboveground live biomass of intact Amazonian rainforests (area 5.76 × 10⁶ km² in 2000) to be 93±23 Pg C, taking into account lianas and small trees. Including dead biomass and belowground biomass would increase this value by approximately 10% and 21%, respectively, but the spatial variation of these additional terms still needs to be quantified.     

Temperature explains global variation in biomass among humid old‐growth forests

Aim To develop and test a simple climate‐based ecophysiological model of above‐ground biomass – an approach that can be applied directly to predicting the effects of climate change on forest carbon stores. Location Humid lowland forests world‐wide. Methods We developed a new approach to modelling the aboveground biomass of old‐growth forest (AGB_max) based on the influences of temperature on gross primary productivity (GPP) and what we call total maintenance cost (TMC), which includes autotrophic respiration as well as leaf, stem and other plant construction required to maintain biomass. We parameterized the models with measured carbon fluxes and tested them by comparing predicted AGB_max with measured AGB for another 109 old‐growth sites. Results Our models explained 57% of the variation in GPP across 95 sites and 79% of the variation in TMC across 17 sites. According to the best‐fit models, the ratio of GPP to maintenance cost per unit biomass (MCB) peaks at 16.5 °C, indicating that this is the air temperature leading to the highest possible AGB_max when temperatures are constant. Seasonal temperature variation generally reduces predicted AGB_max, and thus maritime temperate climates are predicted to have the highest AGB_max. The shift in temperatures from temperate maritime to tropical climates increases MCB more than GPP, and thus decreases AGB_max. Overall, our model explains exactly 50% of the variation in AGB among humid lowland old‐growth forests. Main conclusions Temperature plays an important role in explaining global variation in biomass among humid lowland old‐growth forests, a role that can be understood in terms of the dual effects of temperature on GPP and TMC. Our simple model captures these influences, and could be an important tool for predicting the effects of climate change on forest carbon stores.     

Large altitudinal increase in tree root/shoot ratio in tropical mountain forests of Ecuador

Tropical rain forests decrease in tree height and aboveground biomass (AGB) with increasing elevation. The causes of this phenomenon remain insufficiently understood despite a number of explanations proposed including direct or indirect effects of low temperature on carbon acquisition and carbon investment, adverse soil conditions and impaired nutrient supply. For analysing altitudinal patterns of aboveground/belowground carbon partitioning, we measured fine (<2 mm in diameter) and coarse root (2–5 mm) biomass and necromass and leaf area index (LAI), and estimated AGB from stand structural parameters in five tropical mountain rain forests at 1050, 1540, 1890, 2380 and 3060 m along an altitudinal transect in the South Ecuadorian Andes. Average tree height and AGB were reduced to less than 50% between 1050 and 3060 m, LAI decreased from 5.1 to 2.9. The leaf area reduction must have resulted in a lowered canopy carbon gain and thus may partly explain the reduced tree growth in the high-elevation stands. In contrast, both fine and coarse root biomass significantly increased with elevation across this transect. The ratio of root biomass (fine and coarse) to AGB increased more than ten-fold from 0.04 at 1050 m to 0.43 at 3060 m. Under the assumption that fine root biomass does reflect root productivity, our data indicate a marked belowground shift in C allocation with increasing elevation. Possible explanations for this allocation shift are discussed including reduced N supply due to low temperatures, water logging or adverse soil chemical conditions. We conclude that the fine root system and its activity may hold the key for understanding the impressive reduction in tree size along tropical mountain slopes in Ecuador. Analyses of fine root turnover and longevity in relation to environmental factors along altitudinal transects in tropical mountains are urgently needed.     

Environmental correlates of tree biomass, basal area, wood specific gravity and stem density gradients in Borneo's tropical forests

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. Methods We 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. Results We 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 conclusions Unlike 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.     

Disentangling stand and environmental correlates of aboveground biomass in Amazonian forests

Tropical forests contain an important proportion of the carbon stored in terrestrial vegetation, but estimated aboveground biomass (AGB) in tropical forests varies two‐fold, with little consensus on the relative importance of climate, soil and forest structure in explaining spatial patterns. Here, we present analyses from a plot network designed to examine differences among contrasting forest habitats (terra firme, seasonally flooded, and white‐sand forests) that span the gradient of climate and soil conditions of the Amazon basin. We installed 0.5‐ha plots in 74 sites representing the three lowland forest habitats in both Loreto, Peru and French Guiana, and we integrated data describing climate, soil physical and chemical characteristics and stand variables, including local measures of wood specific gravity (WSG). We use a hierarchical model to separate the contributions of stand variables from climate and soil variables in explaining spatial variation in AGB. AGB differed among both habitats and regions, varying from 78 Mg ha^?1 in white‐sand forest in Peru to 605 Mg ha^?1 in terra firme clay forest of French Guiana. Stand variables including tree size and basal area, and to a lesser extent WSG, were strong predictors of spatial variation in AGB. In contrast, soil and climate variables explained little overall variation in AGB, though they did co‐vary to a limited extent with stand parameters that explained AGB. Our results suggest that positive feedbacks in forest structure and turnover control AGB in Amazonian forests, with richer soils (Peruvian terra firme and all seasonally flooded habitats) supporting smaller trees with lower wood density and moderate soils (French Guianan terra firme) supporting many larger trees with high wood density. The weak direct relationships we observed between soil and climate variables and AGB suggest that the most appropriate approaches to landscape scale modeling of AGB in the Amazon would be based on remote sensing methods to map stand structure.     

Contribution of Dead Wood to Biomass and Carbon Stocks in the Caribbean: St. John, U.S. Virgin Islands

Dead wood is a substantial carbon stock in terrestrial forest ecosystems and hence a critical component of global carbon cycles. Given the limited amounts of dead wood biomass and carbon stock information for Caribbean forests, our objectives were to: (1) describe the relative contribution of down woody materials (DWM) to carbon stocks on the island of St. John; (2) compare these contributions among differing stand characteristics in subtropical moist and dry forests; and (3) compare down woody material carbon stocks on St. John to those observed in other tropical and temperate forests. Our results indicate that dead wood and litter comprise an average of 20 percent of total carbon stocks on St. John in both moist and dry forest life zones. Island-wide, dead wood biomass on the ground ranged from 4.55 to 28.11 Mg/ha. Coarse woody material biomass and carbon content were higher in moist forests than in dry forests. No other down woody material components differed between life zones or among vegetation categories ( P > 0.05). Live tree density was positively correlated with fine woody material and litter in the moist forest life zone ( R = 0.57 and 0.84, respectively) and snag basal area was positively correlated with total down woody material amounts ( R = 0.50) in dry forest. Our study indicates that DWM are important contributors to the total biomass and, therefore, carbon budgets in subtropical systems, and that contributions of DWM on St. John appear to be comparable to values given for similar dry forest systems.     

How drought events during the last century have impacted biomass carbon in Amazonian rainforests

During the last two decades, inventory data show that droughts have reduced biomass carbon sink of the Amazon forest by causing mortality to exceed growth. However, process-based models have struggled to include drought-induced responses of growth and mortality and have not been evaluated against plot data. A process-based model, ORCHIDEE-CAN-NHA, including forest demography with tree cohorts, plant hydraulic architecture and drought-induced tree mortality, was applied over Amazonia rainforests forced by gridded climate fields and rising CO₂ from 1901 to 2019. The model reproduced the decelerating signal of net carbon sink and drought sensitivity of aboveground biomass (AGB) growth and mortality observed at forest plots across selected Amazon intact forests for 2005 and 2010. We predicted a larger mortality rate and a more negative sensitivity of the net carbon sink during the 2015/16 El Niño compared with the former droughts. 2015/16 was indeed the most severe drought since 1901 regarding both AGB loss and area experiencing a severe carbon loss. We found that even if climate change did increase mortality, elevated CO₂ contributed to balance the biomass mortality, since CO₂-induced stomatal closure reduces transpiration, thus, offsets increased transpiration from CO₂-induced higher foliage area.