The Amazon Basin has experienced more variable climate over the last decade, with a severe and widespread drought in 2005 causing large basin‐wide losses of biomass. A drought of similar climatological magnitude occurred again in 2010; however, there has been no basin‐wide ground‐based evaluation of effects on vegetation. We examine to what extent the 2010 drought affected forest dynamics using ground‐based observations of mortality and growth from an extensive forest plot network. We find that during the 2010 drought interval, forests did not gain biomass (net change: −0.43 Mg ha−1, confidence interval (CI): −1.11, 0.19, n = 97), regardless of whether forests experienced precipitation deficit anomalies. This contrasted with a long‐term biomass sink during the baseline pre‐2010 drought period (1998 to pre‐2010) of 1.33 Mg ha−1 yr−1 (CI: 0.90, 1.74, p < 0.01). The resulting net impact of the 2010 drought (i.e., reversal of the baseline net sink) was −1.95 Mg ha−1 yr−1 (CI:−2.77, −1.18; p < 0.001). This net biomass impact was driven by an increase in biomass mortality (1.45 Mg ha−1 yr−1 CI: 0.66, 2.25, p < 0.001) and a decline in biomass productivity (−0.50 Mg ha−1 yr−1, CI:−0.78, −0.31; p < 0.001). Surprisingly, the magnitude of the losses through tree mortality was unrelated to estimated local precipitation anomalies and was independent of estimated local pre‐2010 drought history. Thus, there was no evidence that pre‐2010 droughts compounded the effects of the 2010 drought. We detected a systematic basin‐wide impact of the 2010 drought on tree growth rates across Amazonia, which was related to the strength of the moisture deficit. This impact differed from the drought event in 2005 which did not affect productivity. Based on these ground data, live biomass in trees and corresponding estimates of live biomass in lianas and roots, we estimate that intact forests in Amazonia were carbon neutral in 2010 (−0.07 Pg C yr−1 CI:−0.42, 0.23), consistent with results from an independent analysis of airborne estimates of land‐atmospheric fluxes during 2010. Relative to the long‐term mean, the 2010 drought resulted in a reduction in biomass carbon uptake of 1.1 Pg C, compared to 1.6 Pg C for the 2005 event.

Atmospheric carbon dioxide records indicate that the land surface has acted as a strong global carbon sink over recent decades, with a substantial fraction of this sink probably located in the tropics, particularly in the Amazon. Nevertheless, it is unclear how the terrestrial carbon sink will evolve as climate and atmospheric composition continue to change. Here we analyse the historical evolution of the biomass dynamics of the Amazon rainforest over three decades using a distributed network of 321 plots. While this analysis confirms that Amazon forests have acted as a long-term net biomass sink, we find a long-term decreasing trend of carbon accumulation. Rates of net increase in above-ground biomass declined by one-third during the past decade compared to the 1990s. This is a consequence of growth rate increases levelling off recently, while biomass mortality persistently increased throughout, leading to a shortening of carbon residence times. Potential drivers for the mortality increase include greater climate variability, and feedbacks of faster growth on mortality, resulting in shortened tree longevity. The observed decline of the Amazon sink diverges markedly from the recent increase in terrestrial carbon uptake at the global scale, and is contrary to expectations based on models.

Aboveground tropical tree biomass and carbon storage estimates commonly ignore tree height (H). We estimate the effect of incorporating H on tropics-wide forest biomass estimates in 327 plots across four continents using 42 656 H and diameter measurements and harvested trees from 20 sites to answer the following questions:1. What is the best H-model form and geographic unit to include in biomass models to minimise site-level uncertainty in estimates of destructive biomass?2. To what extent does including H estimates derived in (1) reduce uncertainty in biomass estimates across all 327 plots?3. What effect does accounting for H have on plot- and continental-scale forest biomass estimates?The mean relative error in biomass estimates of destructively harvested trees when including H (mean 0.06), was half that when excluding H (mean 0.13). Power- and Weibull-H models provided the greatest reduction in uncertainty, with regional Weibull-H models preferred because they reduce uncertainty in smaller-diameter classes (≤40 cm D) 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 H 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 H 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 H. After accounting for variation in H, 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 H), then applying our regional relationships implies that carbon storage is overestimated by 35 Pg C (31–39 bootstrapped 95% CI) if H is ignored, assuming that the sampled plots are an unbiased statistical representation of all tropical forest in terms of biomass and height factors. Our results show that tree H is an important allometric factor that needs to be included in future forest biomass estimates to reduce error in estimates of tropical carbon stocks and emissions due to deforestation.

Forest structure and dynamics vary across the Amazon Basin in an east-west gradient coincident with variations in soil fertility and geology. This has resulted in the hypothesis that soil fertility may play an important role in explaining Basin-wide variations in forest biomass, growth and stem turnover rates.Soil samples were collected in a total of 59 different forest plots across the Amazon Basin and analysed for exchangeable cations, carbon, nitrogen and pH, with several phosphorus fractions of likely different plant availability also quantified. Physical properties were additionally examined and an index of soil physical quality developed. Bivariate relationships of soil and climatic properties with above-ground wood productivity, stand-level tree turnover rates, above-ground wood biomass and wood density were first examined with multivariate regression models then applied. Both forms of analysis were undertaken with and without considerations regarding the underlying spatial structure of the dataset.Despite the presence of autocorrelated spatial structures complicating many analyses, forest structure and dynamics were found to be strongly and quantitatively related to edaphic as well as climatic conditions. Basin-wide differences in stand-level turnover rates are mostly influenced by soil physical properties with variations in rates of coarse wood production mostly related to soil phosphorus status. Total soil P was a better predictor of wood production rates than any of the fractionated organic-or inorganic-P pools. This suggests that it is not only the immediately available P forms, but probably the entire soil phosphorus pool that is interacting with forest growth on longer timescales.A role for soil potassium in modulating Amazon forest dynamics through its effects on stand-level wood density was also detected. Taking this into account, otherwise enigmatic variations in stand-level biomass across the Basin were then accounted for through the interacting effects of soil physical and chemical properties with climate. A hypothesis of selfmaintaining forest dynamic feedback mechanisms initiated by edaphic conditions is proposed. It is further suggested that this is a major factor determining endogenous disturbance levels, species composition, and forest productivity across the Amazon Basin.

1 ) Precipitation coefficient of variance Δ B due to height (Mg ha -1 )

Introduction Conclusions References Tables Figures

Positive aboveground biomass trends have been reported from old‐growth forests across the Amazon basin and hypothesized to reflect a large‐scale response to exterior forcing. The result could, however, be an artefact due to a sampling bias induced by the nature of forest growth dynamics. Here, we characterize statistically the disturbance process in Amazon old‐growth forests as recorded in 135 forest plots of the RAINFOR network up to 2006, and other independent research programmes, and explore the consequences of sampling artefacts using a data‐based stochastic simulator. Over the observed range of annual aboveground biomass losses, standard statistical tests show that the distribution of biomass losses through mortality follow an exponential or near‐identical Weibull probability distribution and not a power law as assumed by others. The simulator was parameterized using both an exponential disturbance probability distribution as well as a mixed exponential–power law distribution to account for potential large‐scale blowdown events. In both cases, sampling biases turn out to be too small to explain the gains detected by the extended RAINFOR plot network. This result lends further support to the notion that currently observed biomass gains for intact forests across the Amazon are actually occurring over large scales at the current time, presumably as a response to climate change.

This is a major paper, confirming and greatly extending an earlier finding on the relationships between the vitality of rain forest and the biogeochemical features of the soil across the Amazon basin. The authors demonstrate a gradient in biomass, productivity and turnover from very high values in the base rich soils of the western Amazon, to substantially lower values in the (generally) poor soils of the east. There are strong positive correlations with Cation Exchange Capacity, phosphorus, calcium, magnesium and potassium, but not with nitrogen. Whilst this is of no great surprise to many, the effort of assembling and analysing so many measurements over such a wide geograph-

Leaf size influences many aspects of tree function such as rates of transpiration and photosynthesis and, consequently, often varies in a predictable way in response to environmental gradients. The recent development of pan-Amazonian databases based on permanent botanical plots has now made it possible to assess trends in leaf size across environmental gradients in Amazonia. Previous plot-based studies have shown that the community structure of Amazonian trees breaks down into at least two major ecological gradients corresponding with variations in soil fertility (decreasing from southwest to northeast) and length of the dry season (increasing from northwest to south and east). Here we describe the geographic distribution of leaf size categories based on 121 plots distributed across eight South American countries. We find that the Amazon forest is predominantly populated by tree species and individuals in the mesophyll size class (20.25–182.25 cm2). The geographic distribution of species and individuals with large leaves (>20.25 cm2) is complex but is generally characterized by a higher proportion of such trees in the northwest of the region. Spatially corrected regressions reveal weak correlations between the proportion of large-leaved species and metrics of water availability. We also find a significant negative relationship between leaf size and wood density.

Xylem density is a physical property of wood that varies between individuals, species and environments. It reflects the physiological strategies of trees that lead to growth, survival and reproduction. Measurements of branch xylem density, ρx, were made for 1653 trees representing 598 species, sampled from 87 sites across the Amazon basin. Measured values ranged from 218 kg m−3 for a Cordia sagotii (Boraginaceae) from Mountagne de Tortue, French Guiana to 1130 kg m−3 for an Aiouea sp. (Lauraceae) from Caxiuana, Central Pará, Brazil. Analysis of variance showed significant differences in average ρx across regions and sampled plots as well as significant differences between families, genera and species. A partitioning of the total variance in the dataset showed that species identity (family, genera and species) accounted for 33% with environment (geographic location and plot) accounting for an additional 26%; the remaining "residual" variance accounted for 41% of the total variance. Variations in plot means, were, however, not only accountable by differences in species composition because xylem density of the most widely distributed species in our dataset varied systematically from plot to plot. Thus, as well as having a genetic component, branch xylem density is a plastic trait that, for any given species, varies according to where the tree is growing in a predictable manner. Within the analysed taxa, exceptions to this general rule seem to be pioneer species belonging for example to the Urticaceae whose branch xylem density is more constrained than most species sampled in this study. These patterns of variation of branch xylem density across Amazonia suggest a large functional diversity amongst Amazonian trees which is not well understood.

Abstract. We analysed 1040 individual trees, located in 62 plots across the Amazon Basin for leaf mass per unit area (MA), foliar carbon isotopic composition (δ13C) and leaf level concentrations of C, N, P, Ca, Mg, K and Al. All trees were identified to the species level with the dataset containing 58 families, 236 genera and 508 species, distributed across a wide range of soil types and precipitation regimes. Some foliar characteristics such as MA, [C], [N] and [Mg] emerge as highly constrained by the taxonomic affiliation of tree species, but with others such as [P], [K], [Ca] and δ13C also strongly influenced by site growing conditions. By removing the environmental contribution to trait variation, we find that intrinsic values of most trait pairs coordinate, although different species (characterised by different trait suites) are found at discrete locations along a common axis of coordination. Species that tend to occupy higher fertility soils are characterised by a lower MA and have a higher intrinsic [N], [P], [K], [Mg] and δ13C than their lower fertility counterparts. Despite this consistency, different scaling patterns were observed between low and high fertility sites. Inter-relationships are thus substantially modified by growth environment. Analysing the environmental component of trait variation, we found soil fertility to be the most important predictor, influencing all leaf nutrient concentrations and δ13C and reducing MA. Mean annual temperature was negatively associated with leaf level [N], [P] and [K] concentrations. Total annual precipitation positively influences MA, [C] and δ13C, but with a negative impact on [Mg]. These results provide a first basis for understanding the relationship between the physiological functioning and distribution of tree species across Amazonia.

The net primary productivity (NPP) of tropical forests is one of the most important and least quantified components of the global carbon cycle. Most relevant studies have focused particularly on the quantification of the above-ground coarse wood productivity, and little is known about the carbon fluxes involved in other elements of the NPP, the partitioning of total NPP between its above- and below-ground components and the main environmental drivers of these patterns. In this study we quantify the above- and below-ground NPP of ten Amazonian forests to address two questions: (1) How do Amazonian forests allocate productivity among its above- and below-ground components? (2) How do soil and leaf nutrient status and soil texture affect the productivity of Amazonian forests? Using a standardized methodology to measure the major elements of productivity, we show that NPP varies between 9.3±1.3 Mg C ha−1 yr−1 (mean±standard error), at a white sand plot, and 17.0±1.4 Mg C ha−1 yr−1 at a very fertile Terra Preta site, with an overall average of 12.8±0.9 Mg C ha−1 yr−1. The studied forests allocate on average 64±3% and 36±3% of the total NPP to the above- and below-ground components, respectively. The ratio of above-ground and below-ground NPP is almost invariant with total NPP. Litterfall and fine root production both increase with total NPP, while stem production shows no overall trend. Total NPP tends to increase with soil phosphorus and leaf nitrogen status. However, allocation of NPP to below-ground shows no relationship to soil fertility, but appears to decrease with the increase of soil clay content.

Xylem density is a physical property of wood that varies between individuals, species and environments. It reflects the physiological strategies of trees that lead to growth, survival and reproduction. Measurements of branch xylem density,ρx , were made for 1653 trees representing 598 species, sampled from 87 sites across the Amazon basin. Measured values ranged from 218 kg m −3 for aCordia sagotii (Boraginaceae) from Mountagne de Tortue, French Guiana to 1130 kg m−3 for anAioueasp. (Lauraceae) from Caxiuana, Central Par ́ , Brazil. Analysis of variance showed significant differences in average ρx across regions and sampled plots as well as significant differences between families, genera and species. A partitioning of the total variance in the dataset showed that species identity (family, genera and species) accounted for 33% with environment (geographic location and plot) accounting for an additional 26%; the remaining “residual” variance accounted for 41% of the total variance. Variations in plot means, were, however, not only accountable by differences in species composition because xylem density of the most widely distributed species in our dataset varied systematically from plot to plot. Thus, as well as having a genetic component, branch xylem density is a plastic trait that, for any given species, varies according to where the tree is growing in a predictable manner. Within the analysed taxa, exceptions to this general rule seem to be pioneer species belonging for example to the Urticaceaewhose branch xylem density is more constrained than most species sampled in this study. These patterns of variation of branch xylem density across Amazonia suggest a large functional diversity amongst Amazonian trees which is not well understood.

Abstract. Landscape and environmental variables such as topography, geomorphology, soil types, and climate are important factors affecting forest composition, structure, productivity, and biomass. Here, we combine a network of forest inventories with recently developed global data products from satellite observations in modeling the potential distributions of forest structure and productivity in Amazonia and examine how geomorphology, soil, and precipitation control these distributions. We use the RAINFOR network of forest plots distributed in lowland forests across Amazonia, and satellite observations of tree cover, leaf area index, phenology, moisture, and topographical variations. A maximum entropy estimation (Maxent) model is employed to predict the spatial distribution of several key forest structure parameters: basal area, fraction of large trees, fraction of palms, wood density, productivity, and above-ground biomass at 5 km spatial resolution. A series of statistical tests at selected thresholds as well as across all thresholds and jackknife analysis are used to examine the accuracy of distribution maps and the relative contributions of environmental variables. The final maps were interpreted using soil, precipitation, and geomorphological features of Amazonia and it was found that the length of dry season played a key role in impacting the distribution of all forest variables except the wood density. Soil type had a significant impact on the wood productivity. Most high productivity forests were distributed either on less infertile soils of western Amazonia and Andean foothills, on crystalline shields, and younger alluvial deposits. Areas of low elevation and high density of small rivers of Central Amazonia showed distinct features, hosting mainly forests with low productivity and smaller trees.

Abstract. Leaves in tropical forests come in an enormous variety of sizes and shapes, each of which can be ultimately viewed as an adaptation to the complex problem of optimising the capture of light for photosynthesis. However, the fact that many different shape "strategies" coexist within a habitat demonstrate that there are many other intrinsic and extrinsic factors involved, such as the differential investment in support tissues required for different leaf lamina shapes. Here, we take a macrogeographic approach to understanding the function of different lamina shape categories. Specifically, we use 106 permanent plots spread across the Amazon rainforest basin to: 1) describe the geographic distribution of some simple metrics of lamina shape in plots from across Amazonia, and; 2) identify and quantify relationships between key environmental parameters and lamina shape in tropical forests. Because the plots are not randomly distributed across the study area, achieving this latter objective requires the use of statistics that can account for spatial auto-correlation. We found that between 60–70% of the 2791 species and 83 908 individual trees in the dataset could be classified as having elliptic leaves (= the widest part of the leaf is on an axis in the middle fifth of the long axis of the leaf). Furthermore, the average Amazonian tree leaf is 2.5 times longer than it is wide and has an entire margin. Contrary to theoretical expectations we found little support for the hypothesis that narrow leaves are an adaptation to dry conditions. However, we did find strong regional patterns in leaf lamina length-width ratios and several significant correlations with precipitation variables suggesting that water availability may be exerting an as yet unrecognised selective pressure on leaf shape of rainforest trees. Some support was found for the hypothesis that narrow leaves are an adaptation to low nutrient soils. Furthermore, we found a strong correlation between the proportion of trees with non-entire laminas (dissected, toothed, etc.) and mean annual temperature once again supporting the well documented association that provides a basis for reconstructing past temperature regimes.

Measurements of branch xylem density, Dx, were made for 1466 trees representing 503 species, sampled from 80 sites across the Amazon basin. Measured values ranged from 240 kg m?3 for a Brosimum parinarioides from Tapajos in West Para, Brazil to 1130 kg m?3 for an Aiouea sp. from Caxiuana, Central Para, Brazil. Analysis of variance showed significant differences in average Dx across the sample plots as well as significant differences between families, genera and species. A partitioning of the total variance in the dataset showed that geographic location and plot accounted for 33% of the variation with species identity accounting for an additional 27%; the remaining residual 40% of the variance accounted for by tree to tree (within species) variation. Variations in plot means, were, however, hardly accountable at all by differences in species composition. Rather, it would seem that variations of xylem density at plot level must be explained by the effects of soils and/or climate. This conclusion is supported by the observation that the xylem density of the more widely distributed species varied systematically from plot to plot. Thus, as well as having a genetic component branch xylem density is a plastic trait that, for any given species, varies according to where the tree is growing and in a predictable manner. Exceptions to this general rule may be some pioneers belonging to Pourouma and Miconia and some species within the genera Brosimum, Rinorea and Trichillia which seem to be more constrained in terms of this plasticity than most species sampled as part of this study.

Measurements of branch xylem density, D x , were made for 1466 trees representing 503 species, sampled from 80 sites across the Amazon basin. Measured values ranged from 240 kg m ?3 for a Brosimum parinarioides from Tapajos in West Para, Brazil to 1130 kg m ?3 for an Aiouea sp. from Caxiuana, Central Para, Brazil. Analysis of variance showed significant differences in average D x across the sample plots as well as significant differences between families, genera and species. A partitioning of the total variance in the dataset showed that geographic location and plot accounted for 33% of the variation with species identity accounting for an additional 27%; the remaining "residual" 40% of the variance accounted for by tree to tree (within species) variation. Variations in plot means, were, however, hardly accountable at all by differences in species composition. Rather, it would seem that variations of xylem density at plot level must be explained by the effects of soils and/or climate. This conclusion is supported by the observation that the xylem density of the more widely distributed species varied systematically from plot to plot. Thus, as well as having a genetic component branch xylem density is a plastic trait that, for any given species, varies according to where the tree is growing and in a predictable manner. Exceptions to this general rule may be some pioneers belonging to Pourouma and Miconia and some species within the genera Brosimum, Rinorea and Trichillia which seem to be more constrained in terms of this plasticity than most species sampled as part of this study.