Níro Higuchi

Amazon forests are a key but poorly understood component of the global carbon cycle. If, as anticipated, they dry this century, they might accelerate climate change through carbon losses and changed surface energy balances. We used records from multiple long-term monitoring plots across Amazonia to assess forest responses to the intense 2005 drought, a possible analog of future events. Affected forest lost biomass, reversing a large long-term carbon sink, with the greatest impacts observed where the dry season was unusually intense. Relative to pre-2005 conditions, forest subjected to a 100-millimeter increase in water deficit lost 5.3 megagrams of aboveground biomass of carbon per hectare. The drought had a total biomass carbon impact of 1.2 to 1.6 petagrams (1.2 x 10(15) to 1.6 x 10(15) grams). Amazon forests therefore appear vulnerable to increasing moisture stress, with the potential for large carbon losses to exert feedback on climate change.

The role of the world's forests as a "sink" for atmospheric carbon dioxide is the subject of active debate. Long-term monitoring of plots in mature humid tropical forests concentrated in South America revealed that biomass gain by tree growth exceeded losses from tree death in 38 of 50 Neotropical sites. These forest plots have accumulated 0.71 ton, plus or minus 0.34 ton, of carbon per hectare per year in recent decades. The data suggest that Neotropical forests may be a significant carbon sink, reducing the rate of increase in atmospheric carbon dioxide.

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.

Abstract 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 × 10 6 km 2 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.

Abstract. 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 self-maintaining 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.

*The rich ecology of tropical forests is intimately tied to their moisture status. Multi-site syntheses can provide a macro-scale view of these linkages and their susceptibility to changing climates. Here, we report pan-tropical and regional-scale analyses of tree vulnerability to drought. *We assembled available data on tropical forest tree stem mortality before, during, and after recent drought events, from 119 monitoring plots in 10 countries concentrated in Amazonia and Borneo. *In most sites, larger trees are disproportionately at risk. At least within Amazonia, low wood density trees are also at greater risk of drought-associated mortality, independent of size. For comparable drought intensities, trees in Borneo are more vulnerable than trees in the Amazon. There is some evidence for lagged impacts of drought, with mortality rates remaining elevated 2 yr after the meteorological event is over. *These findings indicate that repeated droughts would shift the functional composition of tropical forests toward smaller, denser-wooded trees. At very high drought intensities, the linear relationship between tree mortality and moisture stress apparently breaks down, suggesting the existence of moisture stress thresholds beyond which some tropical forests would suffer catastrophic tree mortality.

Abstract The net primary production of tropical forests and its partitioning between long‐lived carbon pools (wood) and shorter‐lived pools (leaves, fine roots) are of considerable importance in the global carbon cycle. However, these terms have only been studied at a handful of field sites, and with no consistent calculation methodology. Here we calculate above‐ground coarse wood carbon productivity for 104 forest plots in lowland New World humid tropical forests, using a consistent calculation methodology that incorporates corrections for spatial variations in tree‐size distributions and wood density, and for census interval length. Mean wood density is found to be lower in more productive forests. We estimate that above‐ground coarse wood productivity varies by more than a factor of three (between 1.5 and 5.5 Mg C ha −1 a −1 ) across the Neotropical plots, with a mean value of 3.1 Mg C ha −1 a −1 . There appear to be no obvious relationships between wood productivity and rainfall, dry season length or sunshine, but there is some hint of increased productivity at lower temperatures. There is, however, also strong evidence for a positive relationship between wood productivity and soil fertility. Fertile soils tend to become more common towards the Andes and at slightly higher than average elevations, so the apparent temperature/productivity relationship is probably not a direct one. Coarse wood productivity accounts for only a fraction of overall tropical forest net primary productivity, but the available data indicate that it is approximately proportional to total above‐ground productivity. We speculate that the large variation in wood productivity is unlikely to directly imply an equivalent variation in gross primary production. Instead a shifting balance in carbon allocation between respiration, wood carbon and fine root production seems the more likely explanation.

A previous study by Phillips et al. of changes in the biomass of permanent sample plots in Amazonian forests was used to infer the presence of a regional carbon sink. However, these results generated a vigorous debate about sampling and methodological issues. Therefore we present a new analysis of biomass change in old-growth Amazonian forest plots using updated inventory data. We find that across 59 sites, the above-ground dry biomass in trees that are more than 10 cm in diameter (AGB) has increased since plot establishment by 1.22 +/- 0.43 Mg per hectare per year (ha(-1) yr(-1), where 1 ha = 10(4) m2), or 0.98 +/- 0.38 Mg ha(-1) yr(-1) if individual plot values are weighted by the number of hectare years of monitoring. This significant increase is neither confounded by spatial or temporal variation in wood specific gravity, nor dependent on the allometric equation used to estimate AGB. The conclusion is also robust to uncertainty about diameter measurements for problematic trees: for 34 plots in western Amazon forests a significant increase in AGB is found even with a conservative assumption of zero growth for all trees where diameter measurements were made using optical methods and/or growth rates needed to be estimated following fieldwork. Overall, our results suggest a slightly greater rate of net stand-level change than was reported by Phillips et al. Considering the spatial and temporal scale of sampling and associated studies showing increases in forest growth and stem turnover, the results presented here suggest that the total biomass of these plots has on average increased and that there has been a regional-scale carbon sink in old-growth Amazonian forests during the previous two decades.

Abstract. 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. stand-level 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 amedian −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.

Previous work has shown that tree turnover, tree biomass and large liana densities have increased in mature tropical forest plots in the late twentieth century. These results point to a concerted shift in forest ecological processes that may already be having significant impacts on terrestrial carbon stocks, fluxes and biodiversity. However, the findings have proved controversial, partly because a rather limited number of permanent plots have been monitored for rather short periods. The aim of this paper is to characterize regional-scale patterns of 'tree turnover' (the rate with which trees die and recruit into a population) by using improved datasets now available for Amazonia that span the past 25 years. Specifically, we assess whether concerted changes in turnover are occurring, and if so whether they are general throughout the Amazon or restricted to one region or environmental zone. In addition, we ask whether they are driven by changes in recruitment, mortality or both. We find that: (i) trees 10 cm or more in diameter recruit and die twice as fast on the richer soils of southern and western Amazonia than on the poorer soils of eastern and central Amazonia; (ii) turnover rates have increased throughout Amazonia over the past two decades; (iii) mortality and recruitment rates have both increased significantly in every region and environmental zone, with the exception of mortality in eastern Amazonia; (iv) recruitment rates have consistently exceeded mortality rates; (v) absolute increases in recruitment and mortality rates are greatest in western Amazonian sites; and (vi) mortality appears to be lagging recruitment at regional scales. These spatial patterns and temporal trends are not caused by obvious artefacts in the data or the analyses. The trends cannot be directly driven by a mortality driver (such as increased drought or fragmentation-related death) because the biomass in these forests has simultaneously increased. Our findings therefore indicate that long-acting and widespread environmental changes are stimulating the growth and productivity of Amazon forests.

The loss of tree mass over time from damage can lead to underestimates in above-ground net primary productivity (ANPP) if not accounted for properly. Bias in the allometric relationship between trunk base diameter (Db, at 1.3 m height or above the buttresses) and mass can also lead to systematic errors in ANPP estimates. We developed an unbiased model of the relationship between Db and tree mass using data from 315 trees (≥5 cm Db) harvested in the central Amazon. This model was compared with other theoretical (n=1) and empirical models (n=4). The theoretical model, and one empirical model, made predictions that differed substantially form our central Amazon model. The other three empirical models made predictions that were consistent with our model despite being developed in different tropical forests. Models differed mostly in predicting large tree mass. Using permanent forest inventory plot data, our Db versus tree mass model, and a bole volume model, we estimated that tree damage amounts to 0.9 Mg ha−1 per year (dry mass) of litter production. This damage should be included as a mass loss term when calculating ANPP. Incorporating fine litter data from published studies, we estimated that average ANPP for central Amazon plateau forests is at least 12.9 Mg ha−1 per year (or 6.5 Mg C ha−1 per year). Additional sources of error as described in the text can raise this estimate by as much as 4 Mg ha−1 per year. We hypothesize that tree damage in old-growth forests accounts for a significant portion of age related decline in productivity.

Abstract. 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.

Understanding how tropical forest carbon balance will respond to global change requires knowledge of individual heterotrophic and autotrophic respiratory sources, together with factors that control respiratory variability. We measured leaf, live wood, and soil respiration, along with additional environmental factors over a 1‐yr period in a Central Amazon terra firme forest. Scaling these fluxes to the ecosystem, and combining our data with results from other studies, we estimated an average total ecosystem respiration ( R eco ) of 7.8 μmol·m −2 ·s −1 . Average estimates (per unit ground area) for leaf, wood, soil, total heterotrophic, and total autotrophic respiration were 2.6, 1.1, 3.2, 5.6, and 2.2 μmol·m −2 ·s −1 , respectively. Comparing autotrophic respiration with net primary production (NPP) estimates indicated that only ∼30% of carbon assimilated in photosynthesis was used to construct new tissues, with the remaining 70% being respired back to the atmosphere as autotrophic respiration. This low ecosystem carbon use efficiency (CUE) differs considerably from the relatively constant CUE of ∼0.5 found for temperate forests. Our R eco estimate was comparable to the above‐canopy flux ( F ac ) from eddy covariance during defined sustained high turbulence conditions (when presumably F ac = R eco ) of 8.4 (95% ci = 7.5– 9.4). Multiple regression analysis demonstrated that ∼50% of the nighttime variability in F ac was accounted for by friction velocity ( u *, a measure of turbulence) variables. After accounting for u * variability, mean F ac varied significantly with seasonal and daily changes in precipitation. A seasonal increase in precipitation resulted in a decrease in F ac , similar to our soil respiration response to moisture. The effect of daily changes in precipitation was complex: precipitation after a dry period resulted in a large increase in F ac , whereas additional precipitation after a rainy period had little effect. This response was similar to that of surface litter (coarse and fine), where respiration is greatly reduced when moisture is limiting, but increases markedly and quickly saturates with an increase in moisture.

We used forest inventories, conducted in seventy-two 1-ha permanent plots to assess the variation in tree and palm aboveground live biomass (AGLB), and its relation with soil gradients (texture and nutrients) and topography (altitude and slope). Our plots, located at Reserva Florestal Adolpho Ducke, a 10,000 ha reserve in central Amazonia, near the city of Manaus, were systematically spread over 64 km2. The plots were long (250 m) and narrow (up to 40 m), following elevational contours. Chemical and physical soil analyses were undertaken using topsoil samples, collected in each plot. The plots covered a soil textural gradient ranging from 8% to 98% percent of sand, slopes varying from 0.5° to 27°, and the maximum difference of altitude among plots was 70 m, ranging from 39 to 109 m a.s.l. The mean total AGLB (palm + trees) for stems over 1 cm diameter breast height (dbh = 1.3 m), was 327.8 Mg/ha (ranging from 210.9 to 426.3) and was similar to other studies in the region. We found a two-fold variation in total AGLB estimates among plots, with soil or topography explaining around 20% of this variation. AGLB was positively related to either a soil textural gradient (represented by the percentage of clay) or altitude, but was insensitive to slope. Although slope had no detectable effect on the mean variation of AGLB per plot, it explained 14% of the AGLB in understory and emergent trees. On slopes, AGLB is concentrated mostly in tree classes of small size (1 ≤ dbh < 10 cm), whereas in flat areas, most of the biomass is concentrated in a few big trees. When considering palms and trees separately, we found that tree biomass tended to increase in clay-rich soils (mostly located in high areas), whereas palm biomass was higher in sandy soils (low slopes and bottomlands). This study shows that forests in central Amazonia grow in a heterogeneous environment in relation to soil and topography, and this heterogeneity seems to be in part responsible for differences in structure and AGLB accumulation. In the forest around Manaus, topography can be used to predict aboveground live biomass as an alternative to soil variables, with similar predictive power. Topographic variables can be easily assessed using satellite imagery and could improve current estimates of carbon stocks over large areas in the Amazon.

Abstract Most of the planet's diversity is concentrated in the tropics, which includes many regions undergoing rapid climate change. Yet, while climate‐induced biodiversity changes are widely documented elsewhere, few studies have addressed this issue for lowland tropical ecosystems. Here we investigate whether the floristic and functional composition of intact lowland Amazonian forests have been changing by evaluating records from 106 long‐term inventory plots spanning 30 years. We analyse three traits that have been hypothesized to respond to different environmental drivers (increase in moisture stress and atmospheric CO 2 concentrations): maximum tree size, biogeographic water‐deficit affiliation and wood density. Tree communities have become increasingly dominated by large‐statured taxa, but to date there has been no detectable change in mean wood density or water deficit affiliation at the community level, despite most forest plots having experienced an intensification of the dry season. However, among newly recruited trees, dry‐affiliated genera have become more abundant, while the mortality of wet‐affiliated genera has increased in those plots where the dry season has intensified most. Thus, a slow shift to a more dry‐affiliated Amazonia is underway, with changes in compositional dynamics (recruits and mortality) consistent with climate‐change drivers, but yet to significantly impact whole‐community composition. The Amazon observational record suggests that the increase in atmospheric CO 2 is driving a shift within tree communities to large‐statured species and that climate changes to date will impact forest composition, but long generation times of tropical trees mean that biodiversity change is lagging behind climate change.

The accurate mapping of forest carbon stocks is essential for understanding the global carbon cycle, for assessing emissions from deforestation, and for rational land-use planning. Remote sensing (RS) is currently the key tool for this purpose, but RS does not estimate vegetation biomass directly, and thus may miss significant spatial variations in forest structure. We test the stated accuracy of pantropical carbon maps using a large independent field dataset.Tropical forests of the Amazon basin. The permanent archive of the field plot data can be accessed at: http://dx.doi.org/10.5521/FORESTPLOTS.NET/2014_1.Two recent pantropical RS maps of vegetation carbon are compared to a unique ground-plot dataset, involving tree measurements in 413 large inventory plots located in nine countries. The RS maps were compared directly to field plots, and kriging of the field data was used to allow area-based comparisons.The two RS carbon maps fail to capture the main gradient in Amazon forest carbon detected using 413 ground plots, from the densely wooded tall forests of the north-east, to the light-wooded, shorter forests of the south-west. The differences between plots and RS maps far exceed the uncertainties given in these studies, with whole regions over- or under-estimated by > 25%, whereas regional uncertainties for the maps were reported to be < 5%.Pantropical biomass maps are widely used by governments and by projects aiming to reduce deforestation using carbon offsets, but may have significant regional biases. Carbon-mapping techniques must be revised to account for the known ecological variation in tree wood density and allometry to create maps suitable for carbon accounting. The use of single relationships between tree canopy height and above-ground biomass inevitably yields large, spatially correlated errors. This presents a significant challenge to both the forest conservation and remote sensing communities, because neither wood density nor species assemblages can be reliably mapped from space.

Old-growth forest ecosystems comprise a mosaic of patches in different successional stages, with the fraction of the landscape in any particular state relatively constant over large temporal and spatial scales. The size distribution and return frequency of disturbance events, and subsequent recovery processes, determine to a large extent the spatial scale over which this old-growth steady state develops. Here, we characterize this mosaic for a Central Amazon forest by integrating field plot data, remote sensing disturbance probability distribution functions, and individual-based simulation modeling. Results demonstrate that a steady state of patches of varying successional age occurs over a relatively large spatial scale, with important implications for detecting temporal trends on plots that sample a small fraction of the landscape. Long highly significant stochastic runs averaging 1.0 Mg biomass⋅ha(-1)⋅y(-1) were often punctuated by episodic disturbance events, resulting in a sawtooth time series of hectare-scale tree biomass. To maximize the detection of temporal trends for this Central Amazon site (e.g., driven by CO2 fertilization), plots larger than 10 ha would provide the greatest sensitivity. A model-based analysis of fractional mortality across all gap sizes demonstrated that 9.1-16.9% of tree mortality was missing from plot-based approaches, underscoring the need to combine plot and remote-sensing methods for estimating net landscape carbon balance. Old-growth tropical forests can exhibit complex large-scale structure driven by disturbance and recovery cycles, with ecosystem and community attributes of hectare-scale plots exhibiting continuous dynamic departures from a steady-state condition.

Abstract 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 A sian forests (58.3 ± 7.5 m, 95% CI ), followed by forests in A frica (45.1 ± 2.6 m), A merica (35.8 ± 6.0 m) and A ustralia (35.0 ± 7.4 m), and height–diameter relationships varied similarly; for a given diameter, stems were tallest in A sia, followed by A frica, A merica and A ustralia. 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 A sia 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.

Abstract Tropical forests face increasing climate risk 1,2 , yet our ability to predict their response to climate change is limited by poor understanding of their resistance to water stress. Although xylem embolism resistance thresholds (for example, $$\varPsi $$ <mml:math xmlns:mml="http://www.w3.org/1998/Math/MathML"> <mml:mi>Ψ</mml:mi> </mml:math> 50 ) and hydraulic safety margins (for example, HSM 50 ) are important predictors of drought-induced mortality risk 3–5 , little is known about how these vary across Earth’s largest tropical forest. Here, we present a pan-Amazon, fully standardized hydraulic traits dataset and use it to assess regional variation in drought sensitivity and hydraulic trait ability to predict species distributions and long-term forest biomass accumulation. Parameters $$\varPsi $$ <mml:math xmlns:mml="http://www.w3.org/1998/Math/MathML"> <mml:mi>Ψ</mml:mi> </mml:math> 50 and HSM 50 vary markedly across the Amazon and are related to average long-term rainfall characteristics. Both $$\varPsi $$ <mml:math xmlns:mml="http://www.w3.org/1998/Math/MathML"> <mml:mi>Ψ</mml:mi> </mml:math> 50 and HSM 50 influence the biogeographical distribution of Amazon tree species. However, HSM 50 was the only significant predictor of observed decadal-scale changes in forest biomass. Old-growth forests with wide HSM 50 are gaining more biomass than are low HSM 50 forests. We propose that this may be associated with a growth–mortality trade-off whereby trees in forests consisting of fast-growing species take greater hydraulic risks and face greater mortality risk. Moreover, in regions of more pronounced climatic change, we find evidence that forests are losing biomass, suggesting that species in these regions may be operating beyond their hydraulic limits. Continued climate change is likely to further reduce HSM 50 in the Amazon 6,7 , with strong implications for the Amazon carbon sink.

Several widespread changes in the ecology of old-growth tropical forests have recently been documented for the late twentieth century, in particular an increase in stem turnover (pan-tropical), and an increase in above-ground biomass (neotropical). Whether these changes are synchronous and whether changes in growth are also occurring is not known. We analysed stand-level changes within 50 long-term monitoring plots from across South America spanning 1971-2002. We show that: (i) basal area (BA: sum of the cross-sectional areas of all trees in a plot) increased significantly over time (by 0.10 +/- 0.04 m2 ha(-1) yr(-1), mean +/- 95% CI); as did both (ii) stand-level BA growth rates (sum of the increments of BA of surviving trees and BA of new trees that recruited into a plot); and (iii) stand-level BA mortality rates (sum of the cross-sectional areas of all trees that died in a plot). Similar patterns were observed on a per-stem basis: (i) stem density (number of stems per hectare; 1 hectare is 10(4) m2) increased significantly over time (0.94 +/- 0.63 stems ha(-1) yr(-1)); as did both (ii) stem recruitment rates; and (iii) stem mortality rates. In relative terms, the pools of BA and stem density increased by 0.38 +/- 0.15% and 0.18 +/- 0.12% yr(-1), respectively. The fluxes into and out of these pools-stand-level BA growth, stand-level BA mortality, stem recruitment and stem mortality rates-increased, in relative terms, by an order of magnitude more. The gain terms (BA growth, stem recruitment) consistently exceeded the loss terms (BA loss, stem mortality) throughout the period, suggesting that whatever process is driving these changes was already acting before the plot network was established. Large long-term increases in stand-level BA growth and simultaneous increases in stand BA and stem density imply a continent-wide increase in resource availability which is increasing net primary productivity and altering forest dynamics. Continent-wide changes in incoming solar radiation, and increases in atmospheric concentrations of CO2 and air temperatures may have increased resource supply over recent decades, thus causing accelerated growth and increased dynamism across the world's largest tract of tropical forest.

Abstract. The Amazon basin is likely to be increasingly affected by environmental changes: higher temperatures, changes in precipitation, CO 2 fertilization and habitat fragmentation. To examine the important ecological and biogeochemical consequences of these changes, we are developing an international network, RAINFOR, which aims to monitor forest biomass and dynamics across Amazonia in a co‐ordinated fashion in order to understand their relationship to soil and climate. The network will focus on sample plots established by independent researchers, some providing data extending back several decades. We will also conduct rapid transect studies of poorly monitored regions. Field expeditions analysed local soil and plant properties in the first phase (2001–2002). Initial results suggest that the network has the potential to reveal much information on the continental‐scale relations between forest and environment. The network will also serve as a forum for discussion between researchers, with the aim of standardising sampling techniques and methodologies that will enable Amazonian forests to be monitored in a coherent manner in the coming decades.

Long-term monitoring of distributed, multiple plots is the key to quantify macroecological patterns and changes. Here we examine the evidence for concerted changes in the structure, dynamics and composition of old-growth Amazonian forests in the late twentieth century. In the 1980s and 1990s, mature forests gained biomass and underwent accelerated growth and dynamics, all consistent with a widespread, long-acting stimulation of growth. Because growth on average exceeded mortality, intact Amazonian forests have been a carbon sink. In the late twentieth century, biomass of trees of more than 10cm diameter increased by 0.62+/-0.23tCha-1yr-1 averaged across the basin. This implies a carbon sink in Neotropical old-growth forest of at least 0.49+/-0.18PgCyr-1. If other biomass and necromass components are also increased proportionally, then the old-growth forest sink here has been 0.79+/-0.29PgCyr-1, even before allowing for any gains in soil carbon stocks. This is approximately equal to the carbon emissions to the atmosphere by Amazon deforestation. There is also evidence for recent changes in Amazon biodiversity. In the future, the growth response of remaining old-growth mature Amazon forests will saturate, and these ecosystems may switch from sink to source driven by higher respiration (temperature), higher mortality (as outputs equilibrate to the growth inputs and periodic drought) or compositional change (disturbances). Any switch from carbon sink to source would have profound implications for global climate, biodiversity and human welfare, while the documented acceleration of tree growth and mortality may already be affecting the interactions among millions of species.

Summary Mathematical proofs show that rate estimates, for example of mortality and recruitment, will decrease with increasing census interval when obtained from censuses of non‐homogeneous populations. This census interval effect could be confounding or perhaps even driving conclusions from comparative studies involving such rate estimates. We quantify this artefact for tropical forest trees, develop correction methods and re‐assess some previously published conclusions about forest dynamics. Mortality rates of &gt; 50 species at each of seven sites in Africa, Latin America, Asia and Australia were used as subpopulations to simulate stand‐level mortality rates in a heterogeneous population when census intervals varied: all sites showed decreasing stand mortality rates with increasing census interval length. Stand‐level mortality rates from 14 multicensus long‐term forest plots from Africa, Latin America, Asia and Australia also showed that, on average, mortality rates decreased with increasing census interval length. Mortality, recruitment or turnover rates with differing census interval lengths can be compared using the mean rate of decline from the 14 long‐term plots to standardize estimates to a common census length using λ corr = λ × t 0.08 , where λ is the rate and t is time between censuses in years. This simple general correction should reduce the bias associated with census interval variation, where it is unavoidable. Re‐analysis of published results shows that the pan‐tropical increase in stem turnover rates over the late 20th century cannot be attributed to combining data with differing census intervals. In addition, after correction, Old World tropical forests do not have significantly lower turnover rates than New World sites, as previously reported. Our pan‐tropical best estimate adjusted stem turnover rate is 1.81 ± 0.16% a −1 (mean ± 95% CI, n = 65). As differing census intervals affect comparisons of mortality, recruitment and turnover rates, and can lead to erroneous conclusions, standardized field methods, the calculation of local correction factors at sites where adequate data are available, or the use of our general standardizing formula to take account of sample intervals, are to be recommended.

Stable and radiocarbon isotopes were used to investigate the role of soil clay content in the storage and dynamics of soil carbon in tropical forest soils. Organic matter in clay‐rich Oxisols and Ultisols contains at least two distinct components: (1) material with light δ 13 C signatures and turnover times of decades or less; and (2) clay‐associated, 13 C‐enriched, carbon with turnover times of decades at the surface to millennia at depths &gt;20 cm. Soil texture, in this case clay content, exerts a major control on the amount of slowly cycling carbon and therefore influences the storage and dynamics of carbon in tropical forest soils. Soils in primary tropical forest have been proposed as a potentially large sink for anthropogenic carbon. Comparison of carbon stocks in Oxisols sampled near Manaus, Brazil, shows no measurable change in organic carbon stocks over the past 20 years. Simple models estimating the response of soil carbon pools to a sustained 0.5% yr −1 increase in productivity result in C storage rates of 0.09 to 0.13 MgC ha −1 yr −1 in soil organic matter, with additional potential storage of 0.18 to 0.27 MgC ha −1 yr −1 in surface litter and roots. Most storage occurs in organic matter pools with turnover times less than a decade. Export of carbon in dissolved form from upland terra firme Oxisols likely accounts for &lt;0.2 MgC ha −1 yr −1 , but more work is required to assess the export potential for periodically inundated Spodosols.

Tropical forest structural variation across heterogeneous landscapes may control above-ground carbon dynamics. We tested the hypothesis that canopy structure (leaf area and light availability) - remotely estimated from LiDAR - control variation in above-ground coarse wood production (biomass growth). Using a statistical model, these factors predicted biomass growth across tree size classes in forest near Manaus, Brazil. The same statistical model, with no parameterisation change but driven by different observed canopy structure, predicted the higher productivity of a site 500 km east. Gap fraction and a metric of vegetation vertical extent and evenness also predicted biomass gains and losses for one-hectare plots. Despite significant site differences in canopy structure and carbon dynamics, the relation between biomass growth and light fell on a unifying curve. This supported our hypothesis, suggesting that knowledge of canopy structure can explain variation in biomass growth over tropical landscapes and improve understanding of ecosystem function.

Living trees constitute one of the major stocks of carbon in tropical forests. A better understanding of variations in the dynamics and structure of tropical forests is necessary for predicting the potential for these ecosystems to lose or store carbon, and for understanding how they recover from disturbance. Amazonian tropical forests occur over a vast area that encompasses differences in topography, climate, and geologic substrate. We observed large differences in forest structure, biomass, and tree growth rates in permanent plots situated in the eastern (near Santarém, Pará), central (near Manaus, Amazonas) and southwestern (near Rio Branco, Acre) Amazon, which differed in dry season length, as well as other factors. Forests at the two sites experiencing longer dry seasons, near Rio Branco and Santarém, had lower stem frequencies (460 and 466 ha(-1) respectively), less biodiversity (Shannon-Wiener diversity index), and smaller aboveground C stocks (140.6 and 122.1 Mg C ha(-1)) than the Manaus site (626 trees ha(-1), 180.1 Mg C ha(-1)), which had less seasonal variation in rainfall. The forests experiencing longer dry seasons also stored a greater proportion of the total biomass in trees with >50 cm diameter (41-45 vs 30% in Manaus). Rates of annual addition of C to living trees calculated from monthly dendrometer band measurements were 1.9 (Manaus), 2.8 (Santarém), and 2.6 (Rio Branco) Mg C ha(-1) year(-1). At all sites, trees in the 10-30 cm diameter class accounted for the highest proportion of annual growth (38, 55 and 56% in Manaus, Rio Branco and Santarém, respectively). Growth showed marked seasonality, with largest stem diameter increment in the wet season and smallest in the dry season, though this may be confounded by seasonal variation in wood water content. Year-to-year variations in C allocated to stem growth ranged from nearly zero in Rio Branco, to 0.8 Mg C ha(-1) year(-1) in Manaus (40% of annual mean) and 0.9 Mg C ha(-1) year(-1) (33% of annual mean) in Santarém, though this variability showed no significant relation with precipitation among years. Initial estimates of the C balance of live wood including recruitment and mortality as well as growth suggests that live wood biomass is at near steady-state in Manaus, but accumulating at about 1.5 Mg C ha(-1) at the other two sites. The causes of C imbalance in living wood pools in Santarém and Rio Branco sites are unknown, but may be related to previous disturbance at these sites. Based on size distribution and growth rate differences in the three sites, we predict that trees in the Manaus forest have greater mean age (approximately 240 years) than those of the other two forests (approximately 140 years).

Over the past century almost every ecosystem on Earth has come under the influence of changes in atmospheric composition and climate caused by human activity. Tropical forests are among the most productive and extensive ecosystems, and it has been hypothesized that both the dynamics and biomass of apparently undisturbed, old-growth tropical forests have been changing in response to atmospheric changes. Long-term forest sample plots are a critical tool in detecting and monitoring such changes, and our recent analysis of pan-tropical-forest plot data has suggested that the biomass of tropical forests has been increasing, providing a modest negative feedback on the rate of accumulation of atmospheric CO2. However, it has been argued that some of these old forest plot data sets have significant problems in interpretation because of the use of nonstandardized methodologies. In this paper we examine the extent to which potential field methodological errors may bias estimates of total biomass change by detailed examination of tree-by-tree records from up to 120 Neotropical plots to test predictions from theory. Potential positive biases on measurements of biomass change include a bias in site selection, tree deformities introduced by the measurement process, poor methodologies to deal with tree deformities or buttresses, and nonrecording of negative growth increments. We show that, while it is important to improve and standardize methodologies in current and future forest-plot work, any systematic errors introduced by currently identified biases in past studies are small and calculable. We conclude that most tropical-forest plot data are of useful quality, and that the evidence does still weigh conclusively in favor of a recent increase of biomass in old-growth tropical forests.

Quantifying age structure and tree growth rate of Amazonian forests is essential for understanding their role in the carbon cycle. Here, we use radiocarbon dating and direct measurement of diameter increment to document unexpectedly slow growth rates for trees from three locations spanning the Brazilian Amazon basin. Central Amazon trees, averaging only approximately 1 mm/year diameter increment, grow half as fast as those from areas with more seasonal rainfall to the east and west. Slow growth rates mean that trees can attain great ages; across our sites we estimate 17-50% of trees with diameter >10 cm have ages exceeding 300 years. Whereas a few emergent trees that make up a large portion of the biomass grow faster, small trees that are more abundant grow slowly and attain ages of hundreds of years. The mean age of carbon in living trees (60-110 years) is within the range of or slightly longer than the mean residence time calculated from C inventory divided by annual C allocation to wood growth (40-100 years). Faster C turnover is observed in stands with overall higher rates of diameter increment and a larger fraction of the biomass in large, fast-growing trees. As a consequence, forests can recover biomass relatively quickly after disturbance, whereas recovering species composition may take many centuries. Carbon cycle models that apply a single turnover time for carbon in forest biomass do not account for variations in life strategy and therefore may overestimate the carbon sequestration potential of Amazon forests.

Abstract. Forest structure and dynamics have been noted to vary across the Amazon Basin in an east-west gradient in a pattern which coincides 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. To test this hypothesis and assess the importance of edaphic properties in affect forest structure and dynamics, soil and plant samples were collected in a total of 59 different forest plots across the Amazon Basin. Samples were analysed for exchangeable cations, C, N, pH with various P fractions also determined. Physical properties were also examined and an index of soil physical quality developed. Overall, forest structure and dynamics were found to be strongly and quantitatively related to edaphic conditions. Tree turnover rates emerged to be mostly influenced by soil physical properties whereas forest growth rates were mainly related to a measure of available soil phosphorus, although also dependent on rainfall amount and distribution. On the other hand, large scale variations in forest biomass could not be explained by any of the edaphic properties measured, nor by variation in climate. A new hypothesis of self-maintaining forest dynamic feedback mechanisms initiated by edaphic conditions is proposed. It is further suggested that this is a major factor determining forest disturbance levels, species composition and forest productivity on a Basin wide scale.

Climate change is expected to increase the intensity of extreme precipitation events in Amazonia that in turn might produce more forest blowdowns associated with convective storms. Yet quantitative tree mortality associated with convective storms has never been reported across Amazonia, representing an important additional source of carbon to the atmosphere. Here we demonstrate that a single squall line (aligned cluster of convective storm cells) propagating across Amazonia in January, 2005, caused widespread forest tree mortality and may have contributed to the elevated mortality observed that year. Forest plot data demonstrated that the same year represented the second highest mortality rate over a 15‐year annual monitoring interval. Over the Manaus region, disturbed forest patches generated by the squall followed a power‐law distribution (scaling exponent α = 1.48) and produced a mortality of 0.3–0.5 million trees, equivalent to 30% of the observed annual deforestation reported in 2005 over the same area. Basin‐wide, potential tree mortality from this one event was estimated at 542 ± 121 million trees, equivalent to 23% of the mean annual biomass accumulation estimated for these forests. Our results highlight the vulnerability of Amazon trees to wind‐driven mortality associated with convective storms. Storm intensity is expected to increase with a warming climate, which would result in additional tree mortality and carbon release to the atmosphere, with the potential to further warm the climate system.

Abstract Understanding the processes that determine above‐ground biomass ( AGB ) in Amazonian forests is important for predicting the sensitivity of these ecosystems to environmental change and for designing and evaluating dynamic global vegetation models ( DGVM s). AGB is determined by inputs from woody productivity [woody net primary productivity ( NPP )] and the rate at which carbon is lost through tree mortality. Here, we test whether two direct metrics of tree mortality (the absolute rate of woody biomass loss and the rate of stem mortality) and/or woody NPP , control variation in AGB among 167 plots in intact forest across Amazonia. We then compare these relationships and the observed variation in AGB and woody NPP with the predictions of four DGVM s. The observations show that stem mortality rates, rather than absolute rates of woody biomass loss, are the most important predictor of AGB , which is consistent with the importance of stand size structure for determining spatial variation in AGB . The relationship between stem mortality rates and AGB varies among different regions of Amazonia, indicating that variation in wood density and height/diameter relationships also influences AGB . In contrast to previous findings, we find that woody NPP is not correlated with stem mortality rates and is weakly positively correlated with AGB . Across the four models, basin‐wide average AGB is similar to the mean of the observations. However, the models consistently overestimate woody NPP and poorly represent the spatial patterns of both AGB and woody NPP estimated using plot data. In marked contrast to the observations, DGVM s typically show strong positive relationships between woody NPP and AGB . Resolving these differences will require incorporating forest size structure, mechanistic models of stem mortality and variation in functional composition in DGVM s.

The combined use of high technology instruments and appropriate techniques for discriminating tree species is necessary to improve the biodiversity inventory system in tropical countries. The Fourier-Transform Near-Infrared (FT-NIR) Leaf Spectroscopy appears to be a promising tool for plant species discrimination. In this study, we demonstrate an outstanding performance of FT-NIR, extracted from dried whole leaves, to discriminate closely related species of Eschweilera and Corythophora, Lecythidaceae, a major component of Amazonian forests. We obtained 36 spectral readings, from the adaxial and abaxial surfaces of dried leaves, for 159 individuals representing 10 species. Each spectrum consisted of 1557 FT-NIR absorbance values. We compared the rate of correct specimen (individual tree) identification to species for different datasets and discriminant models, in which individual spectrum consisted of different combinations as to the number of variables (all, stepwise selected), different number of reads per specimen (all reads, adaxial, abaxial, randomly selected), and discriminant models (cross-validation, test set validation). The best results indicated 99.4% of correct specimen identification when we used the average of all 36 spectral readings per specimen and stepwise selected variables. The lowest rate was on average 96.6% when a single spectral reading was used per individual tree (randomly sampled over 100 replicates). Overall, the rate of correct species discrimination was always high and insensible to variable selection, to the different datasets, and to the two major validation models we used. These Species Spectral Signature (SSS) provided better results than current DNA barcoding for plant identification in tropical forests, and represents a fast, low-cost sampling technique. Although further tests are required to assess the potential of FT-NIR spectroscopy for plant identification at broader geographical and phylogenetic scales, the results presented in this paper indicate that SSS extracted from herbarium specimens can be a powerful reference to identify specimens, even when lacking reproductive structures, an so of particular interest for forest inventory and management.

Abstract. Repeated long-term censuses have revealed large-scale spatial patterns in Amazon basin forest structure and dynamism, with some forests in the west of the basin having up to a twice as high rate of aboveground biomass production and tree recruitment as forests in the east. Possible causes for this variation could be the climatic and edaphic gradients across the basin and/or the spatial distribution of tree species composition. To help understand causes of this variation a new individual-based model of tropical forest growth, designed to take full advantage of the forest census data available from the Amazonian Forest Inventory Network (RAINFOR), has been developed. The model allows for within-stand variations in tree size distribution and key functional traits and between-stand differences in climate and soil physical and chemical properties. It runs at the stand level with four functional traits – leaf dry mass per area (Ma), leaf nitrogen (NL) and phosphorus (PL) content and wood density (DW) varying from tree to tree – in a way that replicates the observed continua found within each stand. We first applied the model to validate canopy-level water fluxes at three eddy covariance flux measurement sites. For all three sites the canopy-level water fluxes were adequately simulated. We then applied the model at seven plots, where intensive measurements of carbon allocation are available. Tree-by-tree multi-annual growth rates generally agreed well with observations for small trees, but with deviations identified for larger trees. At the stand level, simulations at 40 plots were used to explore the influence of climate and soil nutrient availability on the gross (ΠG) and net (ΠN) primary production rates as well as the carbon use efficiency (CU). Simulated ΠG, ΠN and CU were not associated with temperature. On the other hand, all three measures of stand level productivity were positively related to both mean annual precipitation and soil nutrient status. Sensitivity studies showed a clear importance of an accurate parameterisation of within- and between-stand trait variability on the fidelity of model predictions. For example, when functional tree diversity was not included in the model (i.e. with just a single plant functional type with mean basin-wide trait values) the predictive ability of the model was reduced. This was also the case when basin-wide (as opposed to site-specific) trait distributions were applied within each stand. We conclude that models of tropical forest carbon, energy and water cycling should strive to accurately represent observed variations in functionally important traits across the range of relevant scales.

The carbon sink capacity of tropical forests is substantially affected by tree mortality. However, the main drivers of tropical tree death remain largely unknown. Here we present a pan-Amazonian assessment of how and why trees die, analysing over 120,000 trees representing > 3800 species from 189 long-term RAINFOR forest plots. While tree mortality rates vary greatly Amazon-wide, on average trees are as likely to die standing as they are broken or uprooted-modes of death with different ecological consequences. Species-level growth rate is the single most important predictor of tree death in Amazonia, with faster-growing species being at higher risk. Within species, however, the slowest-growing trees are at greatest risk while the effect of tree size varies across the basin. In the driest Amazonian region species-level bioclimatic distributional patterns also predict the risk of death, suggesting that these forests are experiencing climatic conditions beyond their adaptative limits. These results provide not only a holistic pan-Amazonian picture of tree death but large-scale evidence for the overarching importance of the growth-survival trade-off in driving tropical tree mortality.

The aboveground biomass content of a region can be estimated by either direct or indirect methods. The direct method corresponds to sampling the biomass content by weighing and extrapolating results to larger areas. It is a destructive and very laborious procedure. The indirect method utilizes formulas whose predictor parameters are obtained from forest inventories. Forest inventories are conducted with the purpose of planning exploitation and land use, and the data derived from inventory are frequently not suitable for biomass estimation. Problems with both these methods increase in the Amazon region, where little information is available on forest biomass. The objective of this paper is to compare a number of biomass equations taken from previous studies and establish an appropriate mathematical formula to estimate the vegetation weight of a particular region in the Amazon forest. A 0.2 ha area of `terra firme' forest was chosen for the comparison. This effort was part of a major forest clearing experiment conducted in Tomé Açu, a town located 250 km south of Belém, the capital of the Brazilian state of Pará. The entire biomass in the 0.2 ha area was weighed. A detailed inventory was carried out in the area prior to the cut and then the biomass was estimated with the inventory data inserted in the selected equations. Fourteen different biomass equations taken from previous studies were used. A comparison of the data of real mass and the mass obtained through the application of the fourteen formulas indicated that the two formulas more suitable for the region are given by (a) FW=αDβHγ, where FW is total fresh weight (kg), D the diameter at breast height (cm), H the total height of the tree (m) and α, β, and γ the regression coefficients, equal to 0.026, 1.529, and 1.747, respectively, and (b) FW=αDβ/(1−M), where α and β are 0.465 and 2.202, respectively, and M the moisture content, defined in terms of mass of moisture per unit mass of fresh biomass. The aboveground fresh biomass weight predicted by the above equations were within ±0.6% of the weight determined in the field.

Intact rainforests have a long-term storage capacity.

Usando um banco de dados com 315 árvores, com DAP≥5 cm, foram testados quatro modelos estatísticos - linear, não linear e dois logarítmicos - para estimar a biomassa de árvores em pé. Os dados foram coletados, de forma destrutiva, na região de Manaus, Estado do Amazonas, em um sítio coberto por floresta de terra-fírme sobre platôs de latossolo amarelo. Em diferentes simulações com diferentes intensidades de amostragem, os quatro modelos estimam precisamente a biomassa, sendo que o afastamento entre a média observada e a estimada, em nenhuma ocasião ultrapassou 5%. As equações para estimar a biomassa de árvores individuais em uma parcela fixa, distintamente para árvores com 5≤ DAP&lt;20 cm e com DAP≥20 cm, são mais consistentes do que o uso de uma única equação para estimar, genericamente, todas as árvores com DAP≥5 cm. O modelo logarítmico com apenas uma variável independente, o DAP, apresenta resultados tão consistentes e precisos quanto os modelos que se utilizam também da variável altura total da árvore. Além do modelo estatístico para estimar o peso da massa fresca total de uma árvore, outras informações são apresentadas, estratificadas nos diferentes compartimentos (tronco, galho grosso, galho fino, folhas e, eventualmente, flores e frutos) de uma árvore, como: concentração de água para estimar o peso da massa seca, concentração carbono e a contribuição do peso de cada compartimento no peso total.

Information on diameter increment and growth patterns for individual trees are important tools for forest management primarily to: (i) select tree species for logging; (ii) selecting tree species for protection; (iii) estimate cutting cycles and (iv) to prescribe silvicultural treatments. Most growth and yield studies in tropical moist forests have emphasized only the stand level instead of individual trees. This study dealt with the analysis of individual growth patterns for 272 trees distributed over two transects (East–West and North–South) measuring 20m×2500 m, which were stratified by plateau, slope and "baixio" (lowland areas near small streams), and tree diameter at breast height (DBH) classes (10cm≤DBH<30 cm, 30cm≤DBH<50 cm and DBH≥50 cm). For each tree, a metal "dendrometer" band was fixed to the trunk and growth in circumference was measured with digital calipers. Measurements were carried out for 19 months, from June 1999 to December 2000; for this study, only 12 months of year 2000 were considered. Individual growth pattern varied significantly over time (P=0.00), and slightly (P=0.08) when the interaction months and DBH classes was included; on the other hand, the signal was very weak (P=0.25) when topographical classes were added to the later interaction, and no signal at all (P=0.89) when the interaction between months, diameter and topographical classes were analyzed. Mean annual diameter increment considering all 272 monitored trees was 1.64±0.21 mm per year (95% CI), falling within the range estimated for the Brazilian Amazon region (1.4–2 mm per year).

We developed an individual-based stochastic-empirical model to simulate the carbon dynamics of live and dead trees in a Central Amazon forest near Manaus, Brazil. The model is based on analyses of extensive field studies carried out on permanent forest inventory plots, and syntheses of published studies. New analyses included: (1) growth suppression of small trees, (2) maximum size (trunk base diameter) for 220 tree species, (3) the relationship between growth rate and wood density, and (4) the growth response of surviving trees to catastrophic mortality (from logging). The model simulates a forest inventory plot, and tracks recruitment, growth, and mortality of live trees, decomposition of dead trees (coarse litter), and how these processes vary with changing environmental conditions. Model predictions were tested against aggregated field data, and also compared with independent measurements including maximum tree age and coarse litter standing stocks. Spatial analyses demonstrated that a plot size of ~10 ha was required to accurately measure wood (live and dead) carbon balance. With the model accurately predicting relevant pools and fluxes, a number of model experiments were performed to predict forest carbon balance response to perturbations including: (1) increased productivity due to CO2 fertilization, (2) a single semi-catastrophic (10%) mortality event, (3) increased recruitment and mortality (turnover) rates, and (4) the combined effects of increased turnover, increased tree growth rates, and decreased mean wood density of new recruits. Results demonstrated that carbon accumulation over the past few decades observed on tropical forest inventory plots (~0.5 Mg C ha−1 year−1) is not likely caused by CO2 fertilization. A maximum 25% increase in woody tissue productivity with a doubling of atmospheric CO2 only resulted in an accumulation rate of 0.05 Mg C ha−1 year−1 for the period 1980–2020 for a Central Amazon forest, or an order of magnitude less than observed on the inventory plots. In contrast, model parameterization based on extensive data from a logging experiment demonstrated a rapid increase in tree growth following disturbance, which could be misinterpreted as carbon sequestration if changes in coarse litter stocks were not considered. Combined results demonstrated that predictions of changes in forest carbon balance during the twenty-first century are highly dependent on assumptions of tree response to various perturbations, and underscores the importance of a close coupling of model and field investigations.

The main objective of this paper is to present and discuss the best methods to estimate live above ground biomass in the Atlantic Forest. The methods presented and conclusions are the products of a workshop entitled "Estimation of Biomass and Carbon Stocks: the Case of Atlantic Rain Forest". Aboveground biomass (AGB) in tropical forests is mainly contained in trees. Tree biomass is a function of wood volume, obtained from the diameter and height, architecture and wood density (dry weight per unit volume of fresh wood). It can be quantified by the direct (destructive) or indirect method where the biomass quantification is estimated using mathematical models. The allometric model can be site specific when elaborated to a particular ecosystem or general that can be used in different sites. For the Atlantic Forest, despite the importance of it, there are only two direct measurements of tree biomass, resulting in allometric models specific for this ecosystem. To select one or other of the available models in the literature to estimate AGB it is necessary take into account what is the main question to be answered and the ease with which it is possible to measure the independent variables in the model. Models that present more accurate estimates should be preferred. However, more simple models (those with one independent variable, usually DBH) can be used when the focus is monitoring the variation in carbon storage through the time. Our observations in the Atlantic Forest suggest that pan-tropical relations proposed by Chave et al. (2005) can be confidently used to estimated tree biomass across biomes as long as tree diameter (DBH), height, and wood density are accounted for in the model. In Atlantic Forest, we recommend the quantification of biomass of lianas, bamboo, palms, tree ferns and epiphytes, which are an important component in this ecosystem. This paper is an outcome of the workshop entitled "Estimation of Biomass and Carbon Stocks: the Case of Atlantic Rain Forest", that was conducted at Ubatuba, São Paulo, Brazil, between 4 and 8 December 2006 as part of the Brazilian project "Ombrophylus Dense Forest floristic composition, structure and function at the Núcleos Picinguaba and Santa Virginia of the Serra do Mar State Park", BIOTA Gradiente.

Abstract. 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. Understanding the relationships between plant traits and ecosystem properties at large spatial scales is important for predicting how compositional change will affect carbon cycling in tropical forests. In this study, we examine the relationships between species wood density, maximum height and above-ground, coarse wood production of trees ≥10 cm diameter (CWP) for 60 Amazonian forest plots. Average species maximum height and wood density are lower in Western than Eastern Amazonia and are negatively correlated with CWP. To test the hypothesis that variation in these traits causes the variation in CWP, we generate plot-level estimates of CWP by resampling the full distribution of tree biomass growth rates whilst maintaining the appropriate tree-diameter and functional-trait distributions for each plot. These estimates are then compared with the observed values. Overall, the estimates do not predict the observed, regional-scale pattern of CWP, suggesting that the variation in community-level trait values does not determine variation in coarse wood productivity in Amazonian forests. Instead, the regional gradient in CWP is caused by higher biomass growth rates across all tree types in Western Amazonia. Therefore, the regional gradient in CWP is driven primarily by environmental factors, rather than the particular functional composition of each stand. These results contrast with previous findings for forest biomass, where variation in wood density, associated with variation in species composition, is an important driver of regional-scale patterns in above-ground biomass. Therefore, in tropical forests, above-ground wood productivity may be less sensitive than biomass to compositional change that alters community-level averages of these plant traits.

Precise estimation of biomass at a regional scale is required for evaluating forest carbon stocks throughout the Amazon. We examined six types of allometric models to identify the best estimator of biomass in primary forests (terra firme) in the northwestern sector of the Brazilian Amazon. We also tested six regression models for estimating tree height. We developed each allometric model using measurements of 101 trees excavated in a primary forest distributed along the upper Rio Negro. A simple power function with stem diameter at breast height D as a single variable was selected as the best model for estimating each biomass component, i.e. above-ground total mass AGW, below-ground total mass BGW, and whole individual mass. Among models developed to estimate tree height H from D, we selected a regression model with a coefficient corresponding to an asymptotic height as the best fit. The D–AGW relationship at our study site differed significantly from models developed previously for other regions of the Amazon. We explain this regional variation in part by regional differences in D–H relationships of sample trees. The D–BGW relationship at our site also differed significantly from that in the central Amazon. However, AGW–BGW relationships were consistent between the upper Rio Negro forest and other forests in the central Amazon, in that the BGW–AGW ratio was constant as 0.136 regardless of tree size. On the basis of D-based allometry and census data from 23 plots established in the upper Rio Negro region, we estimated a stand-level total biomass (dry mass) of 252.6 Mg ha−1. This estimate is at least 73% lower than the potential stand biomass for the region previously suggested by several meta-analyses.

Abstract 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.

Canopy gaps created by wind-throw events, or blowdowns, create a complex mosaic of forest patches varying in disturbance intensity and recovery in the Central Amazon. Using field and remote sensing data, we investigated the short-term (four-year) effects of large (>2000 m(2)) blowdown gaps created during a single storm event in January 2005 near Manaus, Brazil, to study (i) how forest structure and composition vary with disturbance gradients and (ii) whether tree diversity is promoted by niche differentiation related to wind-throw events at the landscape scale. In the forest area affected by the blowdown, tree mortality ranged from 0 to 70%, and was highest on plateaus and slopes. Less impacted areas in the region affected by the blowdown had overlapping characteristics with a nearby unaffected forest in tree density (583 ± 46 trees ha(-1)) (mean ± 99% Confidence Interval) and basal area (26.7 ± 2.4 m(2) ha(-1)). Highly impacted areas had tree density and basal area as low as 120 trees ha(-1) and 14.9 m(2) ha(-1), respectively. In general, these structural measures correlated negatively with an index of tree mortality intensity derived from satellite imagery. Four years after the blowdown event, differences in size-distribution, fraction of resprouters, floristic composition and species diversity still correlated with disturbance measures such as tree mortality and gap size. Our results suggest that the gradients of wind disturbance intensity encompassed in large blowdown gaps (>2000 m(2)) promote tree diversity. Specialists for particular disturbance intensities existed along the entire gradient. The existence of species or genera taking an intermediate position between undisturbed and gap specialists led to a peak of rarefied richness and diversity at intermediate disturbance levels. A diverse set of species differing widely in requirements and recruitment strategies forms the initial post-disturbance cohort, thus lending a high resilience towards wind disturbances at the community level.

The rate of above-ground woody biomass production, W P , in some western Amazon forests exceeds those in the east by a factor of 2 or more. Underlying causes may include climate, soil nutrient limitations and species composition. In this modelling paper, we explore the implications of allowing key nutrients such as N and P to constrain the photosynthesis of Amazon forests, and also we examine the relationship between modelled rates of photosynthesis and the observed gradients in W P . We use a model with current understanding of the underpinning biochemical processes as affected by nutrient availability to assess: (i) the degree to which observed spatial variations in foliar [N] and [P] across Amazonia affect stand-level photosynthesis; and (ii) how these variations in forest photosynthetic carbon acquisition relate to the observed geographical patterns of stem growth across the Amazon Basin. We find nutrient availability to exert a strong effect on photosynthetic carbon gain across the Basin and to be a likely important contributor to the observed gradient in W P . Phosphorus emerges as more important than nitrogen in accounting for the observed variations in productivity. Implications of these findings are discussed in the context of future tropical forests under a changing climate.

Abstract Aim To examine variation in the phylogenetic diversity ( PD ) of tree communities across geographical and environmental gradients in Amazonia. Location Two hundred and eighty‐three c . 1 ha forest inventory plots from across Amazonia. Methods We evaluated PD as the total phylogenetic branch length across species in each plot ( PD ss), the mean pairwise phylogenetic distance between species ( MPD ), the mean nearest taxon distance ( MNTD ) and their equivalents standardized for species richness (ses. PD ss, ses. MPD , ses. MNTD ). We compared PD of tree communities growing (1) on substrates of varying geological age; and (2) in environments with varying ecophysiological barriers to growth and survival. Results PD ss is strongly positively correlated with species richness ( SR ), whereas MNTD has a negative correlation. Communities on geologically young‐ and intermediate‐aged substrates (western and central Amazonia respectively) have the highest SR , and therefore the highest PD ss and the lowest MNTD . We find that the youngest and oldest substrates (the latter on the Brazilian and Guiana Shields) have the highest ses. PD ss and ses. MNTD . MPD and ses. MPD are strongly correlated with how evenly taxa are distributed among the three principal angiosperm clades and are both highest in western Amazonia. Meanwhile, seasonally dry tropical forest (SDTF) and forests on white sands have low PD , as evaluated by any metric. Main conclusions High ses. PD ss and ses. MNTD reflect greater lineage diversity in communities. We suggest that high ses. PD ss and ses. MNTD in western Amazonia results from its favourable, easy‐to‐colonize environment, whereas high values in the Brazilian and Guianan Shields may be due to accumulation of lineages over a longer period of time. White‐sand forests and SDTF are dominated by close relatives from fewer lineages, perhaps reflecting ecophysiological barriers that are difficult to surmount evolutionarily. Because MPD and ses. MPD do not reflect lineage diversity per se , we suggest that PD ss, ses. PD ss and ses. MNTD may be the most useful diversity metrics for setting large‐scale conservation priorities.

The Amazon rain forest sustains the world's highest tree diversity, but it remains unclear why some clades of trees are hyperdiverse, whereas others are not. Using dated phylogenies, estimates of current species richness and trait and demographic data from a large network of forest plots, we show that fast demographic traits--short turnover times--are associated with high diversification rates across 51 clades of canopy trees. This relationship is robust to assuming that diversification rates are either constant or decline over time, and occurs in a wide range of Neotropical tree lineages. This finding reveals the crucial role of intrinsic, ecological variation among clades for understanding the origin of the remarkable diversity of Amazonian trees and forests.

The Amazon forest is integral to the global climate system in part because of the high rate of rainfall recycling through tree transpiration and biodiversity (size and species composition). However, the partitioning of precipitation into evaporation, transpiration and runoff, has been quantified at only a few sites. At our study site in the central Amazon, annual rainfall in 2013 was 2302 mm and latent heat flux measurements made using eddy covariance revealed that 1360 mm (59%) was returned to the atmosphere through evaporation and transpiration. Runoff accounted for 41% of the net ecosystem water loss. Combining annual xylem sap flux estimates with total stand sap wood area, we estimated annual stand transpiration rate to be 851 mm (36% of annual rainfall). Emergent canopy trees (diameter >30 cm; average height of 28 m) were responsible for the majority (71%) of the transpired water flux, recycling potentially 26% of the rainfall back to the atmosphere. By difference, we estimate that 510 mm of intercepted rainwater (22% of rainfall) was evaporated directly back to atmosphere from the canopy. Highest stand transpiration rates occurred during the dryer months due to both increased water vapor pressure deficit and the onset of new leaf flush. This study provides further evidence for convergent water use characteristics of tropical trees and highlights the importance of large trees in tropical moist forests. Large trees have been demonstrated to be vulnerable to drought-related mortality, and thus potentially will make up a critical component of the response of tropical forests to climate change.

Abstract Despite orders of magnitude difference in atmospheric reactivity and great diversity in biological functioning, little is known about monoterpene speciation in tropical forests. Here we report vertically resolved ambient air mixing ratios for 12 monoterpenes in a central Amazon rainforest including observations of the highly reactive cis‐ β ‐ocimene (160 ppt), trans‐ β ‐ocimene (79 ppt), and terpinolene (32 ppt) which accounted for an estimated 21% of total monoterpene composition yet 55% of the upper canopy monoterpene ozonolysis rate. All 12 monoterpenes showed a mixing ratio peak in the upper canopy, with three demonstrating subcanopy peaks in 7 of 11 profiles. Leaf level emissions of highly reactive monoterpenes accounted for up to 1.9% of photosynthesis confirming light‐dependent emissions across several Amazon tree genera. These results suggest that highly reactive monoterpenes play important antioxidant roles during photosynthesis in plants and serve as near‐canopy sources of secondary organic aerosol precursors through atmospheric photooxidation via ozonolysis.

Abstract Surface‐to‐atmosphere emissions of dimethyl sulfide (DMS) may impact global climate through the formation of gaseous sulfuric acid, which can yield secondary sulfate aerosols and contribute to new particle formation. While oceans are generally considered the dominant sources of DMS, a shortage of ecosystem observations prevents an accurate analysis of terrestrial DMS sources. Using mass spectrometry, we quantified ambient DMS mixing ratios within and above a primary rainforest ecosystem in the central Amazon Basin in real‐time (2010–2011) and at high vertical resolution (2013–2014). Elevated but highly variable DMS mixing ratios were observed within the canopy, showing clear evidence of a net ecosystem source to the atmosphere during both day and night in both the dry and wet seasons. Periods of high DMS mixing ratios lasting up to 8 h (up to 160 parts per trillion (ppt)) often occurred within the canopy and near the surface during many evenings and nights. Daytime gradients showed mixing ratios (up to 80 ppt) peaking near the top of the canopy as well as near the ground following a rain event. The spatial and temporal distribution of DMS suggests that ambient levels and their potential climatic impacts are dominated by local soil and plant emissions. A soil source was confirmed by measurements of DMS emission fluxes from Amazon soils as a function of temperature and soil moisture. Furthermore, light‐ and temperature‐dependent DMS emissions were measured from seven tropical tree species. Our study has important implications for understanding terrestrial DMS sources and their role in coupled land‐atmosphere climate feedbacks.

Tree mortality is a key driver of forest community composition and carbon dynamics. Strong winds associated with severe convective storms are dominant natural drivers of tree mortality in the Amazon. Why forests vary with respect to their vulnerability to wind events and how the predicted increase in storm events might affect forest ecosystems within the Amazon are not well understood. We found that windthrows are common in the Amazon region extending from northwest (Peru, Colombia, Venezuela, and west Brazil) to central Brazil, with the highest occurrence of windthrows in the northwest Amazon. More frequent winds, produced by more frequent severe convective systems, in combination with well-known processes that limit the anchoring of trees in the soil, help to explain the higher vulnerability of the northwest Amazon forests to winds. Projected increases in the frequency and intensity of convective storms in the Amazon have the potential to increase wind-related tree mortality. A forest demographic model calibrated for the northwestern and the central Amazon showed that northwestern forests are more resilient to increased wind-related tree mortality than forests in the central Amazon. Our study emphasizes the importance of including wind-related tree mortality in model simulations for reliable predictions of the future of tropical forests and their effects on the Earth' system.

Tropical forests absorb large amounts of atmospheric CO2 through photosynthesis but elevated temperatures suppress this absorption and promote monoterpene emissions. Using 13 CO2 labeling, here we show that monoterpene emissions from tropical leaves derive from recent photosynthesis and demonstrate distinct temperature optima for five groups (Groups 1-5), potentially corresponding to different enzymatic temperature-dependent reaction mechanisms within β-ocimene synthases. As diurnal and seasonal leaf temperatures increased during the Amazonian 2015 El Niño event, leaf and landscape monoterpene emissions showed strong linear enrichments of β-ocimenes (+4.4% °C-1 ) at the expense of other monoterpene isomers. The observed inverse temperature response of α-pinene (-0.8% °C-1 ), typically assumed to be the dominant monoterpene with moderate reactivity, was not accurately simulated by current global emission models. Given that β-ocimenes are highly reactive with respect to both atmospheric and biological oxidants, the results suggest that highly reactive β-ocimenes may play important roles in the thermotolerance of photosynthesis by functioning as effective antioxidants within plants and as efficient atmospheric precursors of secondary organic aerosols. Thus, monoterpene composition may represent a new sensitive 'thermometer' of leaf oxidative stress and atmospheric reactivity, and therefore a new tool in future studies of warming impacts on tropical biosphere-atmosphere carbon-cycle feedbacks.

Abstract Amazon forests account for ~25% of global land biomass and tropical tree species. In these forests, windthrows (i.e., snapped and uprooted trees) are a major natural disturbance, but the rates and mechanisms of recovery are not known. To provide a predictive framework for understanding the effects of windthrows on forest structure and functional composition (DBH ≥10 cm), we quantified biomass recovery as a function of windthrow severity (i.e., fraction of windthrow tree mortality on Landsat pixels, ranging from 0%–70%) and time since disturbance for terra‐firme forests in the Central Amazon. Forest monitoring allowed insights into the processes and mechanisms driving the net biomass change (i.e., increment minus loss) and shifts in functional composition. Windthrown areas recovering for between 4–27 years had biomass stocks as low as 65.2–91.7 Mg/ha or 23%–38% of those in nearby undisturbed forests (~255.6 Mg/ha, all sites). Even low windthrow severities (4%–20% tree mortality) caused decadal changes in biomass stocks and structure. While rates of biomass increment in recovering vegetation were nearly double (6.3 ± 1.4 Mg ha − 1 year − 1 ) those of undisturbed forests (~3.7 Mg ha − 1 year − 1 ), biomass loss due to post‐windthrow mortality was high (up to −7.5 ± 8.7 Mg ha − 1 year − 1 , 8.5 years since disturbance) and unpredictable. Consequently, recovery to 90% of “pre‐disturbance” biomass takes up to 40 years. Resprouting trees contributed little to biomass recovery. Instead, light‐demanding, low‐density genera (e.g., Cecropia , Inga , Miconia, Pourouma, Tachigali, and Tapirira ) were favored, resulting in substantial post‐windthrow species turnover. Shifts in functional composition demonstrate that windthrows affect the resilience of live tree biomass by favoring soft‐wooded species with shorter life spans that are more vulnerable to future disturbances. As the time required for forests to recover biomass is likely similar to the recurrence interval of windthrows triggering succession, windthrows have the potential to control landscape biomass/carbon dynamics and functional composition in Amazon forests.

Forest biomass is an essential indicator for monitoring the Earth's ecosystems and climate. It is a critical input to greenhouse gas accounting, estimation of carbon losses and forest degradation, assessment of renewable energy potential, and for developing climate change mitigation policies such as REDD+, among others. Wall-to-wall mapping of aboveground biomass (AGB) is now possible with satellite remote sensing (RS). However, RS methods require extant, up-to-date, reliable, representative and comparable in situ data for calibration and validation. Here, we present the Forest Observation System (FOS) initiative, an international cooperation to establish and maintain a global in situ forest biomass database. AGB and canopy height estimates with their associated uncertainties are derived at a 0.25 ha scale from field measurements made in permanent research plots across the world's forests. All plot estimates are geolocated and have a size that allows for direct comparison with many RS measurements. The FOS offers the potential to improve the accuracy of RS-based biomass products while developing new synergies between the RS and ground-based ecosystem research communities.

Summary Amazonian droughts are increasing in frequency and severity. However, little is known about how this may influence species‐specific vulnerability to drought across different ecosystem types. We measured 16 functional traits for 16 congeneric species from six families and eight genera restricted to floodplain, swamp, white‐sand or plateau forests of Central Amazonia. We investigated whether habitat distributions can be explained by species hydraulic strategies, and if habitat specialists differ in their vulnerability to embolism that would make water transport difficult during drought periods. We found strong functional differences among species. Nonflooded species had higher wood specific gravity and lower stomatal density, whereas flooded species had wider vessels, and higher leaf and xylem hydraulic conductivity. The P 50 values (water potential at 50% loss of hydraulic conductivity) of nonflooded species were significantly more negative than flooded species. However, we found no differences in hydraulic safety margin among species, suggesting that all trees may be equally likely to experience hydraulic failure during severe droughts. Water availability imposes a strong selection leading to differentiation of plant hydraulic strategies among species and may underlie patterns of adaptive radiation in many tropical tree genera. Our results have important implications for modeling species distribution and resilience under future climate scenarios.

The forests of Amazonia are among the most biodiverse plant communities on Earth. Given the immediate threats posed by climate and land-use change, an improved understanding of how this extraordinary biodiversity is spatially organized is urgently required to develop effective conservation strategies. Most Amazonian tree species are extremely rare but a few are common across the region. Indeed, just 227 'hyperdominant' species account for >50% of all individuals >10 cm diameter at 1.3 m in height. Yet, the degree to which the phenomenon of hyperdominance is sensitive to tree size, the extent to which the composition of dominant species changes with size class and how evolutionary history constrains tree hyperdominance, all remain unknown. Here, we use a large floristic dataset to show that, while hyperdominance is a universal phenomenon across forest strata, different species dominate the forest understory, midstory and canopy. We further find that, although species belonging to a range of phylogenetically dispersed lineages have become hyperdominant in small size classes, hyperdominants in large size classes are restricted to a few lineages. Our results demonstrate that it is essential to consider all forest strata to understand regional patterns of dominance and composition in Amazonia. More generally, through the lens of 654 hyperdominant species, we outline a tractable pathway for understanding the functioning of half of Amazonian forests across vertical strata and geographical locations.

Abstract Aim Water availability is the major driver of tropical forest structure and dynamics. Most research has focused on the impacts of climatic water availability, whereas remarkably little is known about the influence of water table depth and excess soil water on forest processes. Nevertheless, given that plants take up water from the soil, the impacts of climatic water supply on plants are likely to be modulated by soil water conditions. Location Lowland Amazonian forests. Time period 1971–2019. Methods We used 344 long‐term inventory plots distributed across Amazonia to analyse the effects of long‐term climatic and edaphic water supply on forest functioning. We modelled forest structure and dynamics as a function of climatic, soil‐water and edaphic properties. Results Water supplied by both precipitation and groundwater affects forest structure and dynamics, but in different ways. Forests with a shallow water table (depth &lt;5 m) had 18% less above‐ground woody productivity and 23% less biomass stock than forests with a deep water table. Forests in drier climates (maximum cumulative water deficit &lt; −160 mm) had 21% less productivity and 24% less biomass than those in wetter climates. Productivity was affected by the interaction between climatic water deficit and water table depth. On average, in drier climates the forests with a shallow water table had lower productivity than those with a deep water table, with this difference decreasing within wet climates, where lower productivity was confined to a very shallow water table. Main conclusions We show that the two extremes of water availability (excess and deficit) both reduce productivity in Amazon upland ( terra‐firme ) forests. Biomass and productivity across Amazonia respond not simply to regional climate, but rather to its interaction with water table conditions, exhibiting high local differentiation. Our study disentangles the relative contribution of those factors, helping to improve understanding of the functioning of tropical ecosystems and how they are likely to respond to climate change.

The tension between the large global demand for tropical timber, and ecological sustainability and local socioeconomic development in the fragile natural ecosystems of the Amazon region has challenged many generations. In this case study, carried out in an ecotone forest in the northern Brazilian Amazon, we seek to demonstrate how forest management can became more sustainable through silvicultural prescriptions that reduce forest degradation and soil erosion. We evaluated the effects of the sustainable forest management (SFM) on tree diversity, timber volumes, and carbon stocks. A continuous forest inventory was carried out in nine 1-ha permanent plots (900 subplots of 100 m²), distributed in three treatments with three repetitions: T1 = control without selective logging, T2 = SFM, and T3 = SFM + silvicultural treatment of thinning release by tree girdling and poisoning. SFM was carried out with an average of three commercial timber trees harvested per hectare with DBH (diameter at breast height) ≥ 50 cm, corresponding to a timber volume of 20 ± 6 m3 ha-1, equivalent to the removal of 7 ± 2 Mg C ha-1 (5% of the original C). The formation of clearings (gaps) arising from the falling of exploited trees (0.12 ± 0.09 ha) resulted in greater impacts on the forest structure than the construction of 4 m skidder trails (0.06 ± 0.04 ha), with an average of 7 ± 4 damaged trees ha-1 and 5 ± 3 dead trees ha-1. Although six trees species became locally extinct, tree diversity was little altered, and timber volumes and forest carbon volume stocks remained essentially constant at this average logging intensity. Based on our results, sustainable forest management can become an efficient model for land use in the Amazon when harvesting is carried out using this average logging intensity. However, long-term monitoring studies using permanent plots in ecotone forests in the Brazilian Amazon will still be necessary. Quantifying, evaluating, and reporting impacts related to forest management will enable the formation of an empirical basis to support sustanaible forestry practices and for updating environmental legislation, contributing to local socioeconomic development, and maintaining the environmental services provided globally by tropical forests.

Treefall gaps play important roles in both forest dynamics and species diversity, but variability across the full range of gap sizes has not been reported at a regional scale due to the lack of a consistent methodology for their detection. Here we demonstrate the sensitivity of Landsat data for detecting gaps at the subpixel level in the Manaus region, Central Amazon. Spectral mixture analysis (SMA) on treefall gaps was used to map their occurrence across a 3.4 × 104 km2 landscape using the annual change in non-photosynthetic vegetation (ΔNPV) as the change metric. Thirty randomly selected pixels with a spectral signature of a treefall event (i.e. high ΔNPV) were surveyed in the field. The most frequent single-pixel gap size detected using Landsat was ~ 360 m2, and the severity of the disturbance (ΔNPV) exhibited a significant (r2 = 0.32, p = 0.001) correlation with the number of dead trees (> 10 cm diameter at breast height), enabling quantification of the number of downed trees in each gap. To place the importance of these single-pixel disturbances into a broader context, the cumulative disturbance of these gaps was equivalent to 40% of the calculated deforestation across the Manaus region in 2008. Most detected single-pixel gaps consisted of six to eight downed trees covering an estimated area of 250–900 m2. These results highlight the quantitative importance of small blowdowns that have been overlooked in previous satellite remote sensing studies.

Lineages tend to retain ecological characteristics of their ancestors through time. However, for some traits, selection during evolutionary history may have also played a role in determining trait values. To address the relative importance of these processes requires large-scale quantification of traits and evolutionary relationships among species. The Amazonian tree flora comprises a high diversity of angiosperm lineages and species with widely differing life-history characteristics, providing an excellent system to investigate the combined influences of evolutionary heritage and selection in determining trait variation. We used trait data related to the major axes of life-history variation among tropical trees (e.g. growth and mortality rates) from 577 inventory plots in closed-canopy forest, mapped onto a phylogenetic hypothesis spanning more than 300 genera including all major angiosperm clades to test for evolutionary constraints on traits. We found significant phylogenetic signal (PS) for all traits, consistent with evolutionarily related genera having more similar characteristics than expected by chance. Although there is also evidence for repeated evolution of pioneer and shade tolerant life-history strategies within independent lineages, the existence of significant PS allows clearer predictions of the links between evolutionary diversity, ecosystem function and the response of tropical forests to global change.

How plants respond physiologically to leaf warming and low water availability may determine how they will perform under future climate change. In 2015-2016, an unprecedented drought occurred across Amazonia with record-breaking high temperatures and low soil moisture, offering a unique opportunity to evaluate the performances of Amazonian trees to a severe climatic event. We quantified the responses of leaf water potential, sap velocity, whole-tree hydraulic conductance (Kwt), turgor loss and xylem embolism, during and after the 2015-2016 El Niño for five canopy-tree species. Leaf/xylem safety margins (SMs), sap velocity and Kwt showed a sharp drop during warm periods. SMs were negatively correlated with vapour pressure deficit, but had no significant relationship with soil water storage. Based on our calculations of canopy stomatal and xylem resistances, the decrease in sap velocity and Kwt was due to a combination of xylem cavitation and stomatal closure. Our results suggest that warm droughts greatly amplify the degree of trees' physiological stress and can lead to mortality. Given the extreme nature of the 2015-2016 El Niño and that temperatures are predicted to increase, this work can serve as a case study of the possible impact climate warming can have on tropical trees.This article is part of a discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

Prolonged drought stress combined with high leaf temperatures can induce programmed leaf senescence involving lipid peroxidation, and the loss of net carbon assimilation during early stages of tree mortality. Periodic droughts are known to induce widespread tree mortality in the Amazon rainforest, but little is known about the role of lipid peroxidation during drought-induced leaf senescence. In this study, we present observations of green leaf volatile (GLV) emissions during membrane peroxidation processes associated with the combined effects of high leaf temperatures and drought-induced leaf senescence from individual detached leaves and a rainforest ecosystem in the central Amazon. Temperature-dependent leaf emissions of volatile terpenoids were observed during the morning, and together with transpiration and net photosynthesis, showed a post-midday depression. This post-midday depression was associated with a stimulation of C₅ and C₆ GLV emissions, which continued to increase throughout the late afternoon in a temperature-independent fashion. During the 2010 drought in the Amazon Basin, which resulted in widespread tree mortality, green leaf volatile emissions (C₆ GLVs) were observed to build up within the forest canopy atmosphere, likely associated with high leaf temperatures and enhanced drought-induced leaf senescence processes. The results suggest that observations of GLVs in the tropical boundary layer could be used as a chemical sensor of reduced ecosystem productivity associated with drought stress.

Newly created forest edges have significant ecophysiological effects on bordering trees. We studied edge effects on microclimate and tree transpiration rates during wet and dry seasons along a 250 m transect spanning the edge of an unpaved road into an old growth tropical lowland forest in the Central Brazilian Amazon. Canopy openness decreased only minimal from the road (3.68%) towards the forest interior (1.69%). Vapor pressure deficit (measured at 2.2 m height above ground) was lower in the forest interior. The edge effect on microclimate penetrated deeper into the forest (>100 m) during the dry season compared to the wet season (<100 m). Overall, sap flux, and therefore transpiration rate, was 54% higher in trees adjacent to the road compared to forest interior trees. Higher transpiration rates at the forest edge can be explained by higher turbulences and energy exchange of the canopy boundary layer and by a shift in species composition to high water using secondary forest species 25 years after the road construction. Similar changes might be expected for other disturbances affecting local relative humidity and in situations that favor plants with water use traits differing from those found in the forest interior.

Higher levels of taxonomic and evolutionary diversity are expected to maximize ecosystem function, yet their relative importance in driving variation in ecosystem function at large scales in diverse forests is unknown. Using 90 inventory plots across intact, lowland, terra firme, Amazonian forests and a new phylogeny including 526 angiosperm genera, we investigated the association between taxonomic and evolutionary metrics of diversity and two key measures of ecosystem function: aboveground wood productivity and biomass storage. While taxonomic and phylogenetic diversity were not important predictors of variation in biomass, both emerged as independent predictors of wood productivity. Amazon forests that contain greater evolutionary diversity and a higher proportion of rare species have higher productivity. While climatic and edaphic variables are together the strongest predictors of productivity, our results show that the evolutionary diversity of tree species in diverse forest stands also influences productivity. As our models accounted for wood density and tree size, they also suggest that additional, unstudied, evolutionarily correlated traits have significant effects on ecosystem function in tropical forests. Overall, our pan-Amazonian analysis shows that greater phylogenetic diversity translates into higher levels of ecosystem function: tropical forest communities with more distantly related taxa have greater wood productivity. Inventory data from 90 lowland Amazonian forest plots and a phylogeny of 526 angiosperm genera were used to show that taxonomic and phylogenetic diversity are both predictive of wood productivity but not of biomass variation.

Abstract Tall trees are key drivers of ecosystem processes in tropical forest, but the controls on the distribution of the very tallest trees remain poorly understood. The recent discovery of grove of giant trees over 80 meters tall in the Amazon forest requires a reevaluation of current thinking. We used high‐resolution airborne laser surveys to measure canopy height across 282,750 ha of old‐growth and second‐growth forests randomly sampling the entire Brazilian Amazon. We investigated how resources and disturbances shape the maximum height distribution across the Brazilian Amazon through the relations between the occurrence of giant trees and environmental factors. Common drivers of height development are fundamentally different from those influencing the occurrence of giant trees. We found that changes in wind and light availability drive giant tree distribution as much as precipitation and temperature, together shaping the forest structure of the Brazilian Amazon. The location of giant trees should be carefully considered by policymakers when identifying important hot spots for the conservation of biodiversity in the Amazon.

In recent years, the study of skid trail recovery processes has gained momentum. In this review, 121 studies on various aspects of skid trail recovery were evaluated to determine when, where, and how the dominant factors that influence the process of recuperation occur. These studies were located proportionally in the following forest biomes: temperate (60%), tropical (31%), and boreal (9%). Research focused mainly on soil physical properties to ascertain if there had been evidence of recovery. The majority of studies of a decade or less after abandonment demonstrated that heavily used skid trails had not recovered. On the contrary, lightly used skid trails did present full recoveries over the same time span. Soil recovery tended to occur in medium- to coarse-textured soils in temperate and boreal forests. Considering all forest biomes, the impacts of compaction persisted at least two to five decades after logging operations. The impacts were evident in diminished tree heights and volumes from trees growing on skid trails. The last 50 years of research indicates that skid trails, globally, do recover from compaction, albeit slowly.

Our objective was to measure the stable carbon isotope composition of leaf tissue and CO 2 released by respiration (δ r ), and to use this information as an estimate of changes in ecosystem isotopic discrimination that occur in response to seasonal and interannual changes in environmental conditions, and land‐use change (forest‐pasture conversion). We made measurements in primary forest and pastures in the Amazon Basin of Brazil. At the Santarém forest site, δ r values showed a seasonal cycle varying from less than −29‰ to approximately −26‰. The observed seasonal change in δ r was correlated with variation in the observed monthly precipitation. In contrast, there was no significant seasonal variation in δ r at the Manaus forest site (average δ r approximately −28‰), consistent with a narrower range of variation in monthly precipitation than occurred in Santarém. Despite substantial (9‰) vertical variation in leaf δ 13 C, the average δ r values observed for all forest sites were similar to the δ 13 C values of the most exposed sun foliage of the dominant tree species. This suggested that the major portion of recently respired carbon dioxide in these forests was metabolized carbohydrate fixed by the sun leaves at the top of the forest canopy. There was no significant seasonal variation observed in the δ 13 C values of leaf organic matter for the forest sites. We sampled in pastures dominated by the C 4 grass, Brachiaria spp., which is planted after forest vegetation has been cleared. The carbon isotope ratio of respired CO 2 in pastures was enriched in 13 C by approximately 10‰ compared to forest ecosystems. A significant temporal change occurred in δ r after the Manaus pasture was burned. Burning removed much of the encroaching C 3 shrub vegetation and so allowed an increased dominance of the C 4 pasture grass, which resulted in higher δ r values.

Results are described for a forest clearing experiment conducted in the forest reserve of the Tropical Silviculture Experimental Station of the National Institute for Research in the Amazon (INPA), located 60 km from Manaus, the capital of the Brazilian state of Amazonas. An area of 1 ha of virgin forest was cut in July 1995 and left to dry until October of the same year. It was burnt using a procedure similar to that followed by native people in the region to prepare the land for cultivation. The fresh biomass content in the test location, 684.8 t ha −1 , was determined using a formula with parameters from forest inventory. The dry biomass content was 401.5 t ha −1 . Combustion completeness, 20.5 %, was estimated by selecting ten 2 × 2 m 2 subareas, 20 trunks with diameter at breast height (DBH) between 10 and 30 cm, and 71 trunks with DBH higher than 30 cm. Their consumption rates by fire were measured. The 2 × 2 m 2 areas were used to determine the combustion completeness of smaller plant components (characteristic diameters lower than 10 cm) and the trunks to determine the efficiency of the medium and large components (characteristic diameters between 10 and 30 cm and larger than 30 cm, respectively). Combustion completeness for small, medium and large components were 88.2%, 4.39%, and 0.43%, respectively. On the basis of the biomass content (684.4 t ha −1 ), the average biomass moisture and carbon contents (41.6% and 47.8%, respectively, the latter on dry basis), and the obtained combustion completeness (20.5%), the average carbon, CO 2 and CO mass rates released to the atmosphere were estimated to be 37.7, 121, and 8.6 t ha −1 , respectively.

The volatile gas isoprene is emitted in teragrams per annum quantities from the terrestrial biosphere and exerts a large effect on atmospheric chemistry. Isoprene is made primarily from recently fixed photosynthate; however, alternate carbon sources play an important role, particularly when photosynthate is limiting. We examined the relative contribution of these alternate carbon sources under changes in light and temperature, the two environmental conditions that have the strongest influence over isoprene emission. Using a novel real-time analytical approach that allowed us to examine dynamic changes in carbon sources, we observed that relative contributions do not change as a function of light intensity. We found that the classical uncoupling of isoprene emission from net photosynthesis at elevated leaf temperatures is associated with an increased contribution of alternate carbon. We also observed a rapid compensatory response where alternate carbon sources compensated for transient decreases in recently fixed carbon during thermal ramping, thereby maintaining overall increases in isoprene production rates at high temperatures. Photorespiration is known to contribute to the decline in net photosynthesis at high leaf temperatures. A reduction in the temperature at which the contribution of alternate carbon sources increased was observed under photorespiratory conditions, while photosynthetic conditions increased this temperature. Feeding [2-(13)C]glycine (a photorespiratory intermediate) stimulated emissions of [(13)C1-5]isoprene and (13)CO2, supporting the possibility that photorespiration can provide an alternate source of carbon for isoprene synthesis. Our observations have important implications for establishing improved mechanistic predictions of isoprene emissions and primary carbon metabolism, particularly under the predicted increases in future global temperatures.

Tree mortality is a critical process in forest ecosystems, as it influences floristic composition, structure, dynamics, carbon storage, and forest nutrient cycling. However, the mechanisms behind tree death in tropical regions are still poorly characterized. This lack of information is mainly because tree mortality data come from long-interval inventories and studies that measured tree death seasonally are scarce. Here we offer novel insights into the power of fine temporal scale observations and we use a natural history approach to understand the processes and mechanism of tree mortality. We monitored tree mortality every month during one year, in 10 ha of terra-firme forest. To determine the pathways of mortality, we considered the state of the tree at the start of the investigation and the pre and post-mortem characteristics. From November 2010 to October 2011, 67 out of 5808 trees died. Despite the 2010 drought, mortality was highly correlated with monthly rainfall (r = 0.85). In total, six pathways of mortality were assessed. Storms were the main cause of mortality, killing 45% of all dead trees, followed by Biotic/abiotic factors accounting for 30% of tree mortality. The high mortality registered in the rainy season was mostly (78%) due to healthy trees dying uprooted or snapped. Finally, we would benefit from studies that assess mortality on a monthly basis and in combination with quantitative long-term data, we can substantially improve our understanding of the mechanisms behind tree death in the tropics.

High descending winds generated by convective storms are a frequent and a major source of tree mortality disturbance events in the Amazon, affecting forest structure and diversity across a variety of scales, and more frequently observed in western and central portions of the basin. Soil texture in the Central Amazon also varies significantly with elevation along a topographic gradient, with decreasing clay content on plateaus, slopes and valleys respectively. In this study we investigated the critical turning moments (Mcrit - rotational force at the moment of tree failure, an indicator of tree stability or wind resistance) of 60 trees, ranging from 19.0 to 41.1 cm in diameter at breast height (DBH) and located in different topographic positions, and for different species, using a cable-winch load-cell system. Our approach used torque as a measure of tree failure to the point of snapping or uprooting. This approach provides a better understanding of the mechanical forces required to topple trees in tropical forests, and will inform models of wind throw disturbance. Across the topographic positions, size controlled variation in Mcrit was quantified for cardeiro (Scleronema mincranthum (Ducke) Ducke), mata-matá (Eschweilera spp.), and a random selection of trees from 19 other species. Our analysis of Mcrit revealed that tree resistance to failure increased with size (DBH and ABG) and differed among species. No effects of topography or failure mode were found for the species either separately or pooled. For the random species, total variance in Mcrit explained by tree size metrics increased from an R2 of 0.49 for DBH alone, to 0.68 when both DBH and stem fresh wood density (SWD) were included in a multiple regression model. This mechanistic approach allows the comparison of tree vulnerability induced by wind damage across ecosystems, and facilitates the use of forest structural information in ecosystem models that include variable resistance of trees to mortality inducing factors. Our results indicate that observed topographic differences in windthrow vulnerability are likely due to elevational differences in wind velocities, rather than by differences in soil-related factors that might effect Mcrit.

Windthrows are a recurrent disturbance in Amazonia and are an important driver of forest dynamics and carbon storage. In this study, we present for the first time the seasonal and interannual variability of windthrows, focusing on Central Amazonia, and discuss the potential meteorological factors associated with this variability. Landsat images over the 1998–2010 time period were used to detect the occurrence of windthrows, which were identified based on their spectral characteristics and shape. Here, we found that windthrows occurred every year but were more frequent between September and February. Organized convective activity associated with multicell storms embedded in mesoscale convective systems, such as northerly squall lines (that move from northeast to southwest) and southerly squall lines (that move from southwest to northeast) can cause windthrows. We also found that southerly squall lines occurred more frequently than their previously reported ~50 year interval. At the interannual scale, we did not find an association between El Niño-Southern Oscillation (ENSO) and windthrows.

This study was conducted in a humid tropical forest located in Brazil within the Amazon Basin on a heavy clay Ferralsol 24 and 30 years after logging to evaluate the recovery process of skid trails. All timber was extracted with a D6 track-type tractor. In total, four skid trails were evaluated, all logged in 1987 and two of them again in 1993. Systematic sampling was applied to the four trails and two controls. The data collected were rut depth, soil bulk density (BD), soil penetration resistance (PR) and seedling and sapling density. Soil physical properties were determined in three layers from 0 to 15 cm in depth, and seedling density was determined from 36 plots of 4m² in size at the soil sample locations. Rut depths were significantly different (p < 0.05) between years, with the greatest depth of 32 cm recorded for the soil in 1993. The results for BD and PR differed from each other only slightly. In the 0–5 cm depth between the ruts both BD and PR were still significantly higher (p < 0.05) than the control for both years, but within the ruts there were no significant differences between the controls of either year. However, in the 5–15 cm depth within the ruts, BD was always significantly higher (p < 0.001) than the control for both years, as well as was PR for the depths 5–10 cm (p < 0.05) and 10–15 (p < 0.001) within the ruts for both years. The soil layers between the ruts in the depth of 10–15 cm for 1993 were significantly different (p < 0.05) for BD and PR, but the means for this same depth were not root limiting for plants for either BD or PR. The evaluation of seedling and sapling density revealed significant differences for the class of saplings greater than 150 cm. Both the 1987 and the 1993 plots had significantly lower numbers per m², 1987 (p < 0.01) and 1993 (p < 0.001), than the control. There were more saplings located between the ruts than in the ruts for both years, but this was not statistically significant. The conclusion of this study is that partial recovery of the disturbed soil had occurred after 24 and 30 years, with the soil surface horizon within the ruts no different than the undisturbed soil of the control. However, the results show that even 30 years is not sufficient for a full recovery of the soil.

Long-term studies of the dynamics of managed forests in tropical regions are lacking. This study aimed to evaluate the dynamics of a tropical forest, over a 25-year period, that was experimentally logged in 1987 and 1988 and submitted to three different cutting intensities. All trees with diameter at breast height (DBH) ≥ 10 cm have been measured annually since 1990. The three logging intensities that were applied were: light (T1)-trees harvested with DBH ≥ 55 cm; medium (T2)-DBH ≥ 50 cm; and heavy (T3)-DBH ≥ 40 cm. Control plots (T0) were also monitored. The highest mean annual mortality rates (1.82% ± 0.38), recruitment rates (2.93% ± 0.77) and diameter increments (0.30 ± 0.02 cm) occurred in the T3 treatment. Shifts in dynamics of the forest were mainly caused by a striking increase in a fast-growing pioneer species and their high mortality rates. The loss in stocking caused by mortality was greater than to that of replacement by recruitment. The results demonstrated that selective logging altered the natural dynamics of the forest through increased: mortality rates, recruitment and growth rates of the residual trees.

Selective logging is one of the most prominent land uses in tropical forests and although it involves harvesting a limited number of trees, the impact on forest structure, composition and aboveground biomass can be significant. Although these impacts are well documented, what is little known is the extent to which selective logging affects tree floristic composition and its recovery process. Understanding how floristic composition is affected by logging activities is essential for determining subsequent cutting cycles, for the maintenance of carbon stocks and for biodiversity conservation. This research investigates the effect of logging on long-term trends on the recovery of species composition in a tropical forest using a unique logging experiment where measurements have been taken annually over a period of 25 years. Changes in 12 long-term 1-hectare (ha) permanent plots were assessed where different selective logging intensities occurred. In the first years after logging, floristic composition differed widely between intact and selectively logged forests, with exploited areas deviating from pre-logged composition. Over time, exploited areas shifted towards the original composition, with more pronounced changes in this trend after ∼13 years. Shifts in floristic composition were caused mainly by a significant increase in light-demanding fast-growing pioneer species and their subsequent continuous high mortality rates after 13 years of the recovery process. In contrast, the control plots showed similar shifts in composition over time, suggesting external factors such as long-term climate changes may be driving these shifts. The results suggest that 25 years after an experimental selective logging has taken place, floristic composition tends to recover closer to the pre-logged status. Thus, in the absence of further human disturbances, experimental selectively logged forests in low to moderate intensities are compatible with biodiversity conservation, at least during the first cycle of exploitation. Reconciling conservation strategies with the recovery of stocks of commercial timber species would be greatly improved by using these results and lead towards more sustainable forest management plans.

Abstract There is large uncertainty whether Amazon forests will remain a carbon sink as atmospheric CO 2 increases. Hence, we simulated an old‐growth tropical forest using six versions of four terrestrial models differing in scale of vegetation structure and representation of biogeochemical (BGC) cycling, all driven with CO 2 forcing from the preindustrial period to 2100. The models were benchmarked against tree inventory and eddy covariance data from a Brazilian site for present‐day predictions. All models predicted positive vegetation growth that outpaced mortality, leading to continual increases in present‐day biomass accumulation. Notably, the two vegetation demographic models (VDMs) (ED2 and ELM‐FATES) always predicted positive stem diameter growth in all size classes. The field data, however, indicated that a quarter of canopy trees didn't grow over the 15‐year period, and while high interannual variation existed, biomass change was near neutral. With a doubling of CO 2 , three of the four models predicted an appreciable biomass sink (0.77 to 1.24 Mg ha −1 year −1 ). ELMv1‐ECA, the only model used here that includes phosphorus constraints, predicted the lowest biomass sink relative to initial biomass stocks (+21%), lower than the other BGC model, CLM5 (+48%). Models projections differed primarily through variations in nutrient constraints, then carbon allocation, initial biomass, and density‐dependent mortality. The VDM's performance was similar or better than the BGC models run in carbon‐only mode, suggesting that nutrient competition in VDMs will improve predictions. We demonstrate that VDMs are comparable to nondemographic (i.e., “big‐leaf”) models but also include finer scale demography and competition that can be evaluated against field observations.

Logging machinery impacts site soil properties mainly through topsoil removal and compaction. The greatest soil disturbance occurs in the logging infrastructure areas: skid trails, roads and log landings. Although landings generally occupy less of the site than skid trails and roads, they suffer severe compaction usually accompanied with complete topsoil removal to level the soil surface for log decking and loading. Few long-term studies have quantified if landing soil properties recover, with none in the humid tropical forests of Amazonia. The aim of this study was to determine if soil bulk density, penetration resistance and fine root biomass in 27-year-old abandoned log landings had returned to a similar state of the adjacent old-growth forest soils. The working hypothesis was that impacts on soil remained despite the regenerated stand of trees present on the landings. Results revealed that the soil physical properties in the log landings were still significantly higher than the controls. Furthermore, fine root biomass was still below the levels presented in the controls. Therefore, in log landings located on very clayey soils in Central Amazonia, soil compaction persists for at least 27 years.

Every year, logging in the world’s largest tropical forest, located within the Amazon biome, continues unabated. Although it is a preferred alternative to deforestation, the residual stand and site are impacted by logging. The objective of this review was to determine and assess the current state of research throughout Amazonia on the subject of logging impacts. To achieve this goal, a systematic approach was utilized to gather, assess and categorize research articles conducted in the Amazon biome over the last decade. Eligibility for inclusion of articles required demonstration of a direct impact from logging operations. A total of 121 articles were determined to meet the eligibility requirements and were included in this review. Articles were subdivided into three environmental categories: forest (n = 85), wildlife (n = 24) and streams (n = 12). The results of this review demonstrated that impacts from logging activities to the forest site were a direct result of the logging cycle (e.g., how often logging occurs) or logging intensity (e.g., how many trees are felled). The impacts to wildlife varied dependent on species, whereas impacts to streams were affected more by the logging system. Overall, research suggested that to attain sustainability and diminish the impacts from logging, a lower logging intensity of 10–15 m3 ha−1 and a longer logging cycle of 40–60 years would be essential for the long-term viability of forest management in Amazonia.

We studied the effect of experimental logging of 4 ha plots on the regeneration of tree species in a forest 90 km north of Manaus, Amazonas, Brazil. Logging resulted in a total reduction in live wood volume of 44–107 m3 ha−1, although only 63% of this volume was felled, and only 43% removed from the plots. The density of established regeneration (trees and shrubs with diameter at breast height ≤10 cm, and height ≥200 cm) was greater in logged plots than in control plots when measured 3 and 7–8 years after logging. Species richness was also significantly higher in logged plots than in controls. We registered 139 species per 1000 stems, 7–8 years after logging, 143 species per 1000 stems, 3 years after logging, and 136 species per 1000 stems in control plots. Overall species composition was significantly affected by the intensity of logging damage in the plots after 7–8 years, and control plots were significantly different from plots logged 3 years previously. However, changes were not great in relation to natural variation within the forest. Most species increased in density after logging (mean=17%), and the number of individuals belonging to species with commercial value on the local market was 15% greater in logged plots than in control plots. The total potential value of the regeneration, based on the value of wood per m3 (when adult) of the individuals, was 23% higher in logged plots than in control plots, though this difference was not statistically significant. Therefore, enrichment planting is not necessary to maintain either the biodiversity, or potential economic value for wood production, of this forest.

Abstract. 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 (&gt;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.

Abstract. Long-term studies using the RAINFOR network of forest plots have generated significant insights into the spatial and temporal dynamics of forest carbon cycling in Amazonia. In this work, we map and explore the landscape context of several major RAINFOR plot clusters using Landsat ETM+ satellite data. In particular, we explore how representative the plots are of their landscape context, and test whether bias in plot location within landscapes may be influencing the regional mean values obtained for important forest biophysical parameters. Specifically, we evaluate whether the regional variations in wood productivity, wood specific density and above ground biomass derived from the RAINFOR network could be driven by systematic and unintentional biases in plot location. Remote sensing data covering 45 field plots were aggregated to generate landscape maps to identify the specific physiognomy of the plots. In the Landsat ETM+ data, it was possible to spectrally differentiate three types of terra firme forest, three types of forests over Paleovarzea geomorphologycal formation, two types of bamboo-dominated forest, palm forest, Heliconia monodominant vegetation, swamp forest, disturbed forests and land use areas. Overall, the plots were generally representative of the forest physiognomies in the landscape in which they are located. Furthermore, the analysis supports the observed regional trends in those important forest parameters. This study demonstrates the utility of landscape scale analysis of forest physiognomies for validating and supporting the finds of plot based studies. Moreover, the more precise geolocation of many key RAINFOR plot clusters achieved during this research provides important contextual information for studies employing the RAINFOR database.

Abstract. Above-ground tropical tree biomass and carbon storage estimates commonly ignore tree height. We estimate the effect of incorporating height (H) on forest biomass estimates using 37 625 concomitant H and diameter measurements (n = 327 plots) and 1816 harvested trees (n = 21 plots) tropics-wide to answer the following questions: 1. For trees of known biomass (from destructive harvests) which H-model form and geographic scale (plot, region, and continent) most reduces biomass estimate uncertainty? 2. How much does including H relationship estimates derived in (1) reduce uncertainty in biomass estimates across 327 plots spanning four continents? 3. What effect does the inclusion of H in biomass estimates have on plot- and continental-scale forest biomass estimates? The mean relative error in biomass estimates of the destructively harvested trees was half (mean 0.06) when including H, compared to excluding H (mean 0.13). The power- and Weibull-H asymptotic model provided the greatest reduction in uncertainty, with the regional Weibull-H model preferred because it reduces uncertainty in smaller-diameter classes that contain the bulk 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 errors are reduced from 41.8 Mg ha−1 (range 6.6 to 112.4) to 8.0 Mg ha−1 (−2.5 to 23.0) when including $H$. For all plots, above-ground live biomass was 52.2±17.3 Mg ha−1 lower when including H estimates (13%), with the greatest reductions in estimated biomass in Brazilian Shield forests and relatively no change in the Guyana Shield, central Africa and southeast Asia. We show fundamentally different stand structure across the four forested tropical continents, which affects biomass reductions due to $H$. African forests store a greater portion of total biomass in large-diameter trees and trees are on average larger in diameter. This contrasts to forests on all other continents where smaller-diameter trees contain the greatest fractions of total biomass. After accounting for variation in $H$, total biomass per hectare is greatest in Australia, the Guyana Shield, and Asia and lowest in W. Africa, W. Amazonia, and the Brazilian Shield (descending order). Thus, if closed canopy tropical forests span 1668 million km2 and store 285 Pg C, then the overestimate is 35 Pg C if H is ignored, and 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 pantropical carbon stocks and emissions due to deforestation.

Abstract. Isoprene (Is) emissions by plants represent a loss of carbon and energy resources leading to the initial hypothesis that fast growing pioneer species in secondary tropical forests allocate carbon primarily to growth at the expense of isoprenoid defenses. In this study, we quantified leaf isoprene and methanol emissions from the abundant pantropical pioneer tree species Vismia guianensis and ambient isoprene concentrations above a diverse secondary forest in the central Amazon. As photosynthetically active radiation (PAR) was varied (0 to 3000 µmol m−2 s−1) under standard leaf temperature (30 °C), isoprene emissions from V. guianensis increased without saturation up to 80 nmol m−2 s−1. A nonlinear increase in isoprene emissions with respect to net photosynthesis (Pn) resulted in the fraction of Pn dedicated to isoprene emissions increasing with light intensity (up to 2 % of Pn). Emission responses to temperature under standard light conditions (PAR of 1000 µmol m−2 s−1) resulted in the classic uncoupling of isoprene emissions (Topt, iso &gt; 40 °C) from net photosynthesis (Topt, Pn = 30.0–32.5 °C) with up to 7 % of Pn emitted as isoprene at 40 °C. Under standard environmental conditions of PAR and leaf temperature, young V. guianensis leaves showed high methanol emissions, low Pn, and low isoprene emissions. In contrast, mature leaves showed high Pn, high isoprene emissions, and low methanol emissions, highlighting the differential control of leaf phenology over methanol and isoprene emissions. High daytime ambient isoprene concentrations (11 ppbv) were observed above a secondary Amazon rainforest, suggesting that isoprene emissions are common among neotropical pioneer species. The results are not consistent with the initial hypothesis and support a functional role of methanol during leaf expansion and the establishment of photosynthetic machinery and a protective role of isoprene for photosynthesis during high temperature extremes regularly experienced in secondary rainforest ecosystems.

Nonstructural carbon (NSC) reserves act as buffers to sustain tree activity during periods when carbon (C) assimilation does not meet C demand, but little is known about their age and accessibility; we designed a controlled girdling experiment in the Amazon to study tree survival on NSC reserves. We used bomb-radiocarbon (14 C) to monitor the time elapsed between C fixation and release ('age' of substrates). We simultaneously monitored how the mobilization of reserve C affected δ13 CO2 . Six ungirdled control trees relied almost exclusively on recent assimilates throughout the 17 months of measurement. The Δ14 C of CO2 emitted from the six girdled stems increased significantly over time after girdling, indicating substantial remobilization of storage NSC fixed up to 13-14 yr previously. This remobilization was not accompanied by a consistent change in observed δ13 CO2 . These trees have access to storage pools integrating C accumulated over more than a decade. Remobilization follows a very clear reverse chronological mobilization with younger reserve pools being mobilized first. The lack of a shift in the δ13 CO2 might indicate a constant contribution of starch hydrolysis to the soluble sugar pool even outside pronounced stress periods (regular mixing).

Abstract Recent research in the central Amazon suggests that wind is a major agent of disturbance, however, a mechanistic understanding of how wind may lead to tree mortality in Amazonian forests remains unclear. Here we estimated wind speeds necessary to topple central Amazon trees by linking both static and dynamic versions of two wind speed estimation methods (four methods total) to field data on tree failure derived from a static winching study. Static versions of these methods assumed invariant wind characteristics as more trees failed, while dynamic versions updated tree spacing, leaf area index and wind profiles progressively after each tree failure. First, we used a profile method which estimates wind force on individual trees by segments. We calculated drag on each segment and converted drag into basal turning moment, and compared the summed turning moments to the critical turning moment measured in the winching study. Estimated critical wind speeds from the static profile method varied greatly, from 10.75 m s−1 to &amp;gt;120.0 m s−1 with a mean of 45.70 m s−1. Critical wind speeds estimated with static approaches decreased with tree size but were not significantly different between two focal genera. Primary drivers of variation in critical wind speed were tree height and crown size. Second, we used the turning moment coefficient method of Hale, S.E., Gardiner, B., Peace, A., Nicoll, B., Taylor, P. and Pizzirani, S. 2015 Comparison and validation of three versions of a forest wind risk model. Environ. Model. Softw.68, 27–41. doi:10.1016/j.envsoft.2015.01.016.; the static version of this method yielded less-variable estimates, ranging from 18.98 to 52.01 m s−1, with a mean of 30.88 m s−1. Notably, the two static methods for estimating critical wind speeds differed in the trees they identified as having the highest and lowest critical wind speeds. Dynamic variants of the above two methods produced greatly reduced ranges in CWS estimates for our study trees, because after the early tree failures, remaining trees were subject to greater wind penetration into the stand and thus greater loading for a given above-canopy wind speed. CWS estimated with dynamic approaches differed significantly between the focal taxa. Nevertheless, both estimates suggest that wind speeds commonly observed during Amazon storms are sufficient to produce widespread tree damage and mortality.

Tropical forests are acknowledged to be the largest global source of isoprene (C5H8) and monoterpenes (C10H16) emissions, with current synthesis studies suggesting few tropical species emit isoprenoids (20–38%) and do so with highly variable emission capacities, including within the same genera. This apparent lack of a clear phylogenetic thread has created difficulties both in linking isoprenoid function with evolution and for the development of accurate biosphere-atmosphere models. Here, we present a systematic emission study of “hyperdominant” tree species in the Amazon Basin. Across 162 individuals, distributed among 25 botanical families and 113 species, isoprenoid emissions were widespread among both early and late successional species (isoprene: 61.9% of the species; monoterpenes: 15.0%; both isoprene and monoterpenes: 9.7%). The hyperdominant species (69) across the top five most abundant genera, which make up about 50% of all individuals in the Basin, had a similar abundance of isoprenoid emitters (isoprene: 63.8%; monoterpenes: 17.4%; both 11.6%). Among the abundant genera, only Pouteria had a low frequency of isoprene emitting species (15.8% of 19 species). In contrast, Protium, Licania, Inga, and Eschweilera were rich in isoprene emitting species (83.3% of 12 species, 61.1% of 18 species, 100% of 8 species, and 100% of 12 species, respectively). Light response curves of individuals in each of the five genera showed light-dependent, photosynthesis-linked emission rates of isoprene and monoterpenes. Importantly, in every genus, we observed species with light-dependent isoprene emissions together with monoterpenes including β-ocimene. These observations support the emerging view of the evolution of isoprene synthases from β-ocimene synthases. Our results have important implications for understanding isoprenoid function-evolution relationships and the development of more accurate Earth System Models.

Tropical rain forests exist in a broad band across the Earth's warm, moist equatorial regions. They are characterized by their great stature, a wide range of life forms (including many trees with buttresses, thick stemmed climbers, and herbaceous epiphytes), and a large number of tree species. Despite the importance of tropical rain forests as a store of carbon, their role in the carbon cycle is not well understood because they are extensive, variable, and generally more difficult to study than other vegetation types. This chapter discusses the progress in understanding the controls on net primary productivity and the related quantity, and the net ecosystem productivity, which requires close collaboration between disciplines. Studies at the leaf and stand scale, using eco physiological and eddy covariance techniques, are advancing one's understanding of the temporal changes. Thereafter, scaling up to whole regions and biomes still requires remotely sensed data on the distribution of land-surface cover, as well as the use of interpolated climatological data from the ground or from global circulation models to drive the models. There is a need to develop new approaches to this difficult problem, perhaps using large-scale experimentation and observation. One aspect of environmental change that has received attention is the influence of forest edges that are created during logging and burning.

As três parcelas permanentes usadas neste estudo são testemunhas (não perturbadas) de um experimento de manejo florestal do Instituto Nacional de Pesquisas da Amazônia, no município de Manaus (AM). Essas parcelas têm sido monitoradas desde 1980, mas para efeito deste estudo, foram consideradas 12 medições repetidas no período 1986-2000. Durante este período, o fenômeno El Niño (seca anormal na região) ocorreu em duas ocasiões, em 1992-93 e 1997-98, sendo que o último foi seguido do La Niña (chuva anormal na região), em 1999. Devido a esses fenômenos, as taxas de recrutamento e mortalidade foram iguais, 0,7%, durante o período observado. No entanto, a acumulação (fixação na árvore) de carbono, foi de 16 toneladas métricas, dando um incremento periódico anual significativo (p = 0,039), em torno de 1,2 t/ha/ano. CARBON BALANCE AND DYNAMICS OF PRIMARY VEGETATION IN THE CENTRAL AMAZON Abstract The three permanent forest inventory plots used for this study were control plots (not disturbed) from a forest management project of the National Institute of Amazon Research (INPA) in the Brazilian State of Amazonas. These plots have been monitored since 1980, although for this study the period from 1986-2000 was considered. During this period, the El Niño phenomenon, which causes increased drought in the region, occurred on two occasions (1992-93 and 1997-98), followed by La Niña which causes increased precipitation in the region (1999-2000). Despite of this change in climate, recruitment and mortality rates were equal throughout the period at 0.7% yr-1. During the same period, carbon accumulation in forest biomass was 16 Mg, resulting in a statistically significant (p = 0.039) increase of about 1.2 Mg biomass ha-1 yr-1.

Ecology Letters (2009) 12: E22–E25 Abstract Lloyd et al. (2009) question the methods, concepts and conclusions of Fisher et al. (2008) . We address these assertions, and place our work into a broader context. We demonstrate the veracity of Fisher et al. , and further show that lack of data for intermediate‐scale tree mortality disturbance events for old‐growth tropical forests might prevent robust extrapolation of forest plot biomass accumulation data, and accurate estimates of distribution parameters such as power‐law exponents ( α ).

ABSTRACT Aim To assess the hypotheses that compound leaves of trees in the Amazon forest are an adaptation to drought and/or rapid growth. Location Amazon rain forest, South America. Methods Genera from 137 permanent forest plots spread across Amazonia were classified into those with compound leaves and those with simple leaves. Metrics of compound leaf prevalence were then calculated for each plot and regression models that accounted for spatial autocorrelation were used to identify associations between climate variables and compound leaf structure. We also tested for associations between compound leaf structure and a variety of ecological variables related to life history and growth strategies, including wood density, annual increase in diameter and maximum height. Results One plant family, Fabaceae, accounts for 53% of compound‐leaved individuals in the dataset, and has a geographical distribution strongly centred on north‐east Amazonia. On exclusion of Fabaceae from the analysis we found no significant support for the seasonal drought hypothesis. However, we found evidence supporting the rapid growth hypothesis, with possession of compound leaves being associated with faster diameter growth rates and lower wood densities. Main conclusion This study provides evidence that possession of compound leaves constitutes one of a suite of traits and life‐history strategies that promote rapid growth in rain forest trees. Our findings highlight the importance of carefully considering the geographical distribution of dominant taxa and spatial clustering of data points when inferring ecological causation from environment–trait associations.

In Amazonian non-flooded forests with a moderate dry season, many trees do not form anatomically definite annual rings. Alternative indicators of annual rings, such as the oxygen (δ(18)Owc) and carbon stable isotope ratios of wood cellulose (δ(13)Cwc), have been proposed; however, their applicability in Amazonian forests remains unclear. We examined seasonal variations in the δ(18)Owc and δ(13)Cwc of three common species (Eschweilera coriacea, Iryanthera coriacea, and Protium hebetatum) in Manaus, Brazil (Central Amazon). E. coriacea was also sampled in two other regions to determine the synchronicity of the isotopic signals among different regions. The annual cyclicity of δ(18)Owc variation was cross-checked by (14)C dating. The δ(18)Owc showed distinct seasonal variations that matched the amplitude observed in the δ(18)O of precipitation, whereas seasonal δ(13)Cwc variations were less distinct in most cases. The δ(18)Owc variation patterns were similar within and between some individual trees in Manaus. However, the δ(18)Owc patterns of E. coriacea differed by region. The ages of some samples estimated from the δ(18)Owc cycles were offset from the ages estimated by (14)C dating. In the case of E. coriacea, this phenomenon suggested that missing or wedging rings may occur frequently even in well-grown individuals. Successful cross-dating may be facilitated by establishing δ(18)Owc master chronologies at both seasonal and inter-annual scales for tree species with distinct annual rings in each region.

Abstract. Windthrows change forest structure and species composition in central Amazon forests. However, the effects of widespread tree mortality associated with wind disturbances on soil properties have not yet been described in this vast region. We investigated short-term effects (7 years after disturbance) of widespread tree mortality caused by a squall line event from mid-January of 2005 on soil carbon stocks and concentrations in a central Amazon terra firme forest. The soil carbon stock (averaged over a 0–30 cm depth profile) in disturbed plots (61.4 ± 8.2 Mg ha−1, mean ±95 % confidence interval) was marginally higher (p = 0.09) than that from undisturbed plots (47.7 ± 13.6 Mg ha−1). The soil organic carbon concentration in disturbed plots (2.0 ± 0.17 %) was significantly higher (p &lt; 0.001) than that from undisturbed plots (1.36 ± 0.24 %). Moreover, soil carbon stocks were positively correlated with soil clay content (r2 = 0.332, r = 0.575 and p = 0.019) and with tree mortality intensity (r2 = 0.257, r = 0.506 and p = 0.045). Our results indicate that large inputs of plant litter associated with large windthrow events cause a short-term increase in soil carbon content, and the degree of increase is related to soil clay content and tree mortality intensity. The higher carbon content and potentially higher nutrient availability in soils from areas recovering from windthrows may favor forest regrowth and increase vegetation resilience.

C₁ metabolism in plants is known to be involved in photorespiration, nitrogen and amino acid metabolism, as well as methylation and biosynthesis of metabolites and biopolymers. Although the flux of carbon through the C₁ pathway is thought to be large, its intermediates are difficult to measure and relatively little is known about this potentially ubiquitous pathway. In this study, we evaluated the C₁ pathway and its integration with the central metabolism using aqueous solutions of 13C-labeled C₁ and C₂ intermediates delivered to branches of the tropical species Inga edulis via the transpiration stream. Delivery of [13C]methanol and [13C]formaldehyde rapidly stimulated leaf emissions of [13C]methanol, [13C]formaldehyde, [13C]formic acid, and 13CO₂, confirming the existence of the C1 pathway and rapid interconversion between methanol and formaldehyde. However, while [13C]formate solutions stimulated emissions of 13CO₂, emissions of [13C]methanol or [13C]formaldehyde were not detected, suggesting that once oxidation to formate occurs it is rapidly oxidized to CO₂ within chloroplasts. 13C-labeling of isoprene, a known photosynthetic product, was linearly related to 13CO₂ across C₁ and C₂ ([13C₂]acetate and [2-13C]glycine) substrates, consistent with reassimilation of C₁, respiratory, and photorespiratory CO₂. Moreover, [13C]methanol and [13C]formaldehyde induced a quantitative labeling of both carbon atoms of acetic acid emissions, possibly through the rapid turnover of the chloroplastic acetyl-CoA pool via glycolate oxidation. The results support a role of the C₁ pathway to provide an alternative carbon source for glycine methylation in photorespiration, enhance CO₂ concentrations within chloroplasts, and produce key C₂ intermediates (e.g., acetyl-CoA) central to anabolic and catabolic metabolism.