Abstract. The Moderate Resolution Imaging Spectroradiometer vegetation continuous fields (MODIS VCF) Earth observation product is widely used to estimate forest cover changes, parameterise vegetation and Earth System models, and as a reference for validation or calibration where field data is limited. However, although limited independent validations of MODIS VCF have shown that MODIS VCF's accuracy decreases when estimating tree cover in sparsely-vegetated areas, such as in tropical savannas, no study has yet assessed the impact this may have on the VCF based tree cover distributions used by many in their research. Using tropical forest and savanna inventory data collected by the TROpical Biomes in Transition (TROBIT) project, we produce a series of corrections that take into account (i) the spatial disparity between the in-situ plot size and the MODIS VCF pixel, and (ii) the trees' spatial distribution within in-situ plots. We then applied our corrections to areas identified as forest or savanna in the International Geosphere-Biosphere Programme (IGBP) land cover mapping product. All IGBP classes identified as savanna show substantial increases in cover after correction, indicating that the most recent version of MODIS VCF consistently underestimates woody cover in tropical savannas. We estimate that MODIS VCF could be underestimating tropical tree cover by between 9–15 %. Models that use VCF as their benchmark could be underestimating the carbon uptake in forest-savanna areas and misrepresenting forest-savanna dynamics. While more detailed in-situ field data is necessary to produce more accurate and reliable corrections, we recommend caution when using MODIS VCF in tropical savannas.
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<p>Fire is an important cause of disturbance in terrestrial ecosystems and can have a major impact on biodiversity. Long-term forest plot monitoring in Mato Grosso state (Brazil) indicates lowest species diversity in plots that have been burned multiple times and increasing species richness with time since the last fire. Furthermore, there is a strong positive relationship between species richness and basal area in unburnt and once-burnt plots, especially in the large tree stratum. We used high-resolution Landsat and PALSAR data at varying spatio-temporal (single and bi-temporal) scales to (a) assess Vegetation Indices sensitive to varying fire severity for different tropical forest species; (b) quantify fire severity and basal area/ biomass changes (1999, 2006, 2010, 2013); and (c) quantify the sensitivity of L-band backscatter to fuel load, moisture content and basal area/ biomass dynamics.</p>

<p>During extreme drought events, aboveground biomass (AGB) dynamics in Amazonian forests are altered through reduced productivity and increased tree mortality and carbon loss. Tree adaptations developed in response to historical drought may reduce the severity of carbon loss. Past droughts were likely associated with fire, which produced Pyrogenic Carbon (PyC), a form of carbon formed by the incomplete combustion of biomass burn and fossil fuel. PyC has specific properties that improve soil fertility and water holding capacity and decrease aluminium toxicity, among others. PyC can be found in different concentrations across the Amazon Basin, since it can be produced by local fires and aerosol deposition. It is unknown whether PyC could explain tree adaptations or contributes to Amazon forest dynamics, especially for extreme drought events. We hypothesize that PyC in soil can serve as a proxy of fire history and fire/drought adaptations and also support the forest during drought events because of its properties, decreasing mortality rates and maintaining rates of AGB gain equivalent to a non-extreme drought year. To evaluate this hypothesis, we used a dataset with more than 70 plots with repeat censuses distributed across the Amazon Basin and classified extreme drought events using maximum cumulative water deficit (MCWD) analysis. Soil samples were collected from the same plots during an intensive fieldwork campaign and PyC was quantified by hydrogen pyrolysis (HyPy). Forest plots were classified into high and low PyC based on the median across the whole dataset. Our preliminary results show that during extreme drought events, plots that have a greater concentration of PyC had significantly higher rates of AGB gain when compared with plots with lower concentrations of PyC (t-test, p < 0.05). During non-extreme drought years there was no significant difference in rates of AGB gain between plots with different concentrations of PyC. When we focus on plots with lower concentrations of PyC there is a significant decrease in rates of AGB gain during drought years compared to non-extreme drought years (t-test, p < 0.05). However, in plots with high concentrations of PyC there is no significant difference in rates of AGB gain, showing trees are able to maintain normal forest dynamics during extreme drought years. We conclude that PyC has an important role in mediating drought resistance and productivity in Amazonian forests.</p>

Abstract. The representation of disturbance is a critical factor in land-surface modelling, but is generally poorly constrained in carbon cycle models. In particular, land-use change and fire can be treated as large-scale disturbances without full representation of their underlying complexities and interactions. Here we describe developments to the land surface model JULES (Joint UK Land Environment Simulator) to represent land-use change and fire as separate disturbances. We use the HYDE (History Database of the Global Environment) land cover dataset to analyse the impact of land-use change on global vegetation, and couple the fire model INFERNO (INteractive Fire and Emission algoRithm for Natural envirOnments) to dynamic vegetation within JULES to assess how the representation of disturbance affects the simulation of present day vegetation. We test model performance, evaluating the inclusion of land use and fire disturbance against standard benchmarks. Using the Manhattan Metric, overall disturbance improves the simulation of vegetation cover compared to observations by up to 53 %. Grasses show an improvement of up to 52 %, with biases in extent reduced from −66 % to 13 %. Total woody cover improves by up to 121 % from a reduction in forest extent in the tropics, although simulated tree cover is now too sparse in some areas. Disturbance generally decreases tree and shrub cover and increases grasses. The results show that the disturbances provide important contributions to the realistic modelling of vegetation on a global scale, although in some areas fire and land-use together result in over-disturbance. This work provides a substantial contribution towards representing the full complexity and interactions between land-use change and fire that could be used in Earth System Models.
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Drought and fire reduce productivity and increase tree mortality in tropical forests. Fires also produce pyrogenic carbon (PyC), which persists in situ for centuries to millennia, and represents a legacy of past fires, potentially improving soil fertility and water holding capacity and selecting for the survival and recruitment of certain tree life-history (or successional) strategies. We investigated whether PyC is correlated with physicochemical soil properties, wood density, aboveground carbon (AGC) dynamics and forest resistance to severe drought. To achieve our aim, we used an Amazon-wide, long-term plot network, in forests without known recent fires, integrating site-specific measures of forest dynamics, soil properties and a unique soil PyC concentration database. We found that forests with higher concentrations of soil PyC had both higher soil fertility and lower wood density. Soil PyC was not associated with AGC dynamics in non-drought years. However, during extreme drought events (10% driest years), forests with higher concentrations of soil PyC experienced lower reductions in AGC gains (woody growth and recruitment), with this drought-immunizing effect increasing with drought severity. Forests with a legacy of ancient fires are therefore more likely to continue to grow and recruit under increased drought severity. Forests with high soil PyC concentrations (third quartile) had 3.8% greater AGC gains under mean drought, but 33.7% greater under the most extreme drought than forests with low soil PyC concentrations (first quartile), offsetting losses of up to 0.68 Mg C ha–1yr–1 of AGC under extreme drought events. This suggests that ancient fires have legacy effects on current forest dynamics, by altering soil fertility and favoring tree species capable of continued growth and recruitment during droughts. Therefore, mature forest that experienced fires centuries or millennia ago may have greater resistance to current short-term droughts.

AbstractTropical forests face increasing climate risk1,2, 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, $$\varPsi $$
. Ψ
. 50) and hydraulic safety margins (for example, HSM50) are important predictors of drought-induced mortality risk3–5, 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 $$\varPsi $$
. Ψ
. 50 and HSM50 vary markedly across the Amazon and are related to average long-term rainfall characteristics. Both $$\varPsi $$
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. 50 and HSM50 influence the biogeographical distribution of Amazon tree species. However, HSM50 was the only significant predictor of observed decadal-scale changes in forest biomass. Old-growth forests with wide HSM50 are gaining more biomass than are low HSM50 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 HSM50 in the Amazon6,7, with strong implications for the Amazon carbon sink.

Aim: to investigate the geographic patterns and ecological correlates in the geographic distribution of the most common tree dispersal modes in Amazonia (endozoochory, synzoochory, anemochory and hydrochory). We examined if the proportional abundance of these dispersal modes could be explained by the availability of dispersal agents (disperser-availability hypothesis) and/or the availability of resources for constructing zoochorous fruits (resource-availability hypothesis). Time period: Tree-inventory plots established between 1934 and 2019. Major taxa studied: Trees with a diameter at breast height (DBH) ≥ 9.55 cm. Location: Amazonia, here defined as the lowland rain forests of the Amazon River basin and the Guiana Shield. Methods: We assigned dispersal modes to a total of 5433 species and morphospecies within 1877 tree-inventory plots across terra-firme, seasonally flooded, and permanently flooded forests. We investigated geographic patterns in the proportional abundance of dispersal modes. We performed an abundance-weighted mean pairwise distance (MPD) test and fit generalized linear models (GLMs) to explain the geographic distribution of dispersal modes. Results: Anemochory was significantly, positively associated with mean annual wind speed, and hydrochory was significantly higher in flooded forests. Dispersal modes did not consistently show significant associations with the availability of resources for constructing zoochorous fruits. A lower dissimilarity in dispersal modes, resulting from a higher dominance of endozoochory, occurred in terra-firme forests (excluding podzols) compared to flooded forests. Main conclusions: the disperser-availability hypothesis was well supported for abiotic dispersal modes (anemochory and hydrochory). The availability of resources for constructing zoochorous fruits seems an unlikely explanation for the distribution of dispersal modes in Amazonia. The association between frugivores and the proportional abundance of zoochory requires further research, as tree recruitment not only depends on dispersal vectors but also on conditions that favour or limit seedling recruitment across forest types.

For monitoring and reporting forest carbon stocks and fluxes, many countries in the tropics and subtropics rely on default values of forest aboveground biomass (AGB) from the Intergovernmental Panel on Climate Change (IPCC) guidelines for National Greenhouse Gas (GHG) Inventories. Default IPCC forest AGB values originated from 2006, and are relatively crude estimates of average values per continent and ecological zone. The 2006 default values were based on limited plot data available at the time, methods for their derivation were not fully clear, and no distinction between successional stages was made. As part of the 2019 Refinement to the 2006 IPCC Guidelines for GHG Inventories, we updated the default AGB values for tropical and subtropical forests based on AGB data from >25 000 plots in natural forests and a global AGB map where no plot data were available. We calculated refined AGB default values per continent, ecological zone, and successional stage, and provided a measure of uncertainty. AGB in tropical and subtropical forests varies by an order of magnitude across continents, ecological zones, and successional stage. Our refined default values generally reflect the climatic gradients in the tropics, with more AGB in wetter areas. AGB is generally higher in old-growth than in secondary forests, and higher in older secondary (regrowth >20 years old and degraded/logged forests) than in young secondary forests (20 years old). While refined default values for tropical old-growth forest are largely similar to the previous 2006 default values, the new default values are 4.0-7.7-fold lower for young secondary forests. Thus, the refined values will strongly alter estimated carbon stocks and fluxes, and emphasize the critical importance of old-growth forest conservation. We provide a reproducible approach to facilitate future refinements and encourage targeted efforts to establish permanent plots in areas with data gaps.

Tree death is a key process for our understanding of how forests are and will respond to global change. The extensive forests across the southern Amazonia edge—the driest, warmest and most fragmented of the Amazon regions—provide a window onto what the future of large parts of Amazonia may look like. Understanding tree mortality and its drivers here is essential to anticipate the process across other parts of the basin. Using 10 years of data from a widespread network of long-term forest plots, we assessed how trees die (standing, broken or uprooted) and used generalised mixed-effect models to explore the contribution of plot-, species- and tree-level factors to the likelihood of tree death. Most trees died from stem breakage (54%); a smaller proportion died standing (41%), while very few were uprooted (5%). The mortality rate for standing dead trees was greatest in forests subject to the most intense dry seasons. While trees with the crown more exposed to light were more prone to death from mechanical damage, trees less exposed were more susceptible to death from drought. At the species level, mortality rates were lowest for those species with the greatest wood density. At the individual tree level, physical damage to the crown via branch breakage was the strongest predictor of tree death. Synthesis. Wind- and water deficit-driven disturbances are the main causes of tree death in southern Amazonia edge which is concerning considering the predicted increase in seasonality for Amazonia, especially at the edge. Tree mortality here is greater than any in other Amazonian region, thus any increase in mortality here may represent a tipping point for these forests.

Brosimum rubescens, a tree species with Neotropical distribution, can achieve local monodominance in Southern Amazonian forests. Understanding how and why this species varies across space and time is important because the monodominance of some species alters ecosystem complexity. Here we evaluated the fundamental ecological niche of B. rubescens by species distribution models (SDM), combining predictive environmental variables with occurrence points, and determined the temporal persistence and how the spatial distribution patterns of this species vary with different environmental predictive variables. To generate the SDMs, we incorporated predictive environmental variables as main components of climatic, hydric and edaphic variables. All algorithms showed higher performance in spatial predictions for hydric variables and for the combination of climatic, hydric and edaphic variables. We identified that the potential niches of B. rubescens seem to be defined by climatic fluctuations, with the edaphic conditions not limiting the presence of this species in the evaluated spatial scale. From the last glacial maximum to the present, this species seems to have increased its spatial amplitude; however, from the present to the future, predictions suggest that B. rubescens will experience a considerable loss of its range. Our findings showed independent and combined effects of different environmental variables, allowing us to identify which are limiting or facilitating the spatial distribution of B. rubescens. We corroborate the spatial persistence and geographical fidelity of the species’ distribution patterns over time. Graphical abstract: [Figure not available: see fulltext.].

Fire has a historical role in tropical forests related to past climate and ancient land use spanning the Holocene; however, it is unclear from charcoal records how fire varied at different spatiotemporal scales and what sampling strategies are required to determine fire history and their effects. We evaluated fire variation in structurally intact, terra-firme Amazon forests, by intensive soil charcoal sampling from three replicate soil pits in sites in Guyana and northern and southern Peru. We used radiocarbon (14C) measurement to assess (1) locally, how the timing of fires represented in our sample varied across the surface of forest plots and with soil depth, (2) basin-wide, how the age of fires varies across climate and environmental gradients, and (3) how many samples are appropriate when applying the 14C approach to assess the date of last fire. Considering all 14C dates (n = 33), the most recent fires occurred at a similar time at each of the three sites (median ages: 728–851 cal years BP), indicating that in terms of fire disturbance at least, these forests could be considered old-growth. The number of unique fire events ranged from 1 to 4 per pit and from 4 to 6 per site. Based upon our sampling strategy, the N-Peru site—with the highest annual precipitation—had the most fire events. Median fire return intervals varied from 455 to 2,950 cal years BP among sites. Based on available dates, at least three samples (1 from the top of each of 3 pits) are required for the sampling to have a reasonable likelihood of capturing the most recent fire for forests with no history of a recent fire. The maximum fire return interval for two sites was shorter than the time since the last fire, suggesting that over the past ∼800 years these forests have undergone a longer fire-free period than the past 2,000–3,500 years. Our analysis from terra-firme forest soils helps to improve understanding of changes in fire regime, information necessary to evaluate post-fire legacies on modern vegetation and soil and to calibrate models to predict forest response to fire under climate change.

Tree diversity and composition in Amazonia are known to be strongly determined by the water supplied by precipitation. Nevertheless, within the same climatic regime, water availability is modulated by local topography and soil characteristics (hereafter referred to as local hydrological conditions), varying from saturated and poorly drained to well-drained and potentially dry areas. While these conditions may be expected to influence species distribution, the impacts of local hydrological conditions on tree diversity and composition remain poorly understood at the whole Amazon basin scale. Using a dataset of 443 1-ha non-flooded forest plots distributed across the basin, we investigate how local hydrological conditions influence 1) tree alpha diversity, 2) the community-weighted wood density mean (CWM-wd) – a proxy for hydraulic resistance and 3) tree species composition. We find that the effect of local hydrological conditions on tree diversity depends on climate, being more evident in wetter forests, where diversity increases towards locations with well-drained soils. CWM-wd increased towards better drained soils in Southern and Western Amazonia. Tree species composition changed along local soil hydrological gradients in Central-Eastern, Western and Southern Amazonia, and those changes were correlated with changes in the mean wood density of plots. Our results suggest that local hydrological gradients filter species, influencing the diversity and composition of Amazonian forests. Overall, this study shows that the effect of local hydrological conditions is pervasive, extending over wide Amazonian regions, and reinforces the importance of accounting for local topography and hydrology to better understand the likely response and resilience of forests to increased frequency of extreme climate events and rising temperatures.

The Moderate Resolution Imaging Spectroradiometer Vegetation Continuous Fields (MODIS VCF) Earth observation product is widely used to estimate forest cover changes and to parameterize vegetation and Earth system models and as a reference for validation or calibration where field data are limited. However, although limited independent validations of MODIS VCF have shown that MODIS VCF's accuracy decreases when estimating tree cover in sparsely vegetated areas such as tropical savannas, no study has yet assessed the impact this may have on the VCF-based tree cover data used by many in their research. Using tropical forest and savanna inventory data collected by the Tropical Biomes in Transition (TROBIT) project, we produce a series of calibration scenarios that take into account (i) the spatial disparity between the in situ plot size and the MODIS VCF pixel and (ii) the trees' spatial distribution within in situ plots. To identify if a disparity also exists in products trained using VCF, we used a similar approach to evaluate the finer-scale Landsat Tree Canopy Cover (TCC) product. For MODIS VCF, we then applied our calibrations to areas identified as forest or savanna in the International Geosphere-Biosphere Programme (IGBP) land cover mapping product. All IGBP classes identified as "savanna"show substantial increases in cover after calibration, indicating that the most recent version of MODIS VCF consistently underestimates woody cover in tropical savannas. We also found that these biases are propagated in the finer-scale Landsat TCC. Our scenarios suggest that MODIS VCF accuracy can vary substantially, with tree cover underestimation ranging from 0% to 29%. Models that use MODIS VCF as their benchmark could therefore be underestimating the carbon uptake in forest-savanna areas and misrepresenting forest-savanna dynamics. Because of the limited in situ plot number, our results are designed to be used as an indicator of where the product is potentially more or less reliable. Until more in situ data are available to produce more accurate calibrations, we recommend caution when using uncalibrated MODIS VCF data in tropical savannas.

Tree mortality rates and the modes of tree death have recently been extensively investigated in the Amazon. However, efforts to describe these processes have not been well distributed across the basin. No study has yet investigated in depth tree mortality process in the unique low, open, bamboo-dominated forests of southwestern Amazonia, a region with a distinct climate and the epicenter of recent severe drought events. Here, we investigated the leading ways that trees die in the terra-firme forests of the southwestern Brazilian Amazon, to understand whether the dynamics of mortality differ from those recorded in other parts of the basin. Using data from six permanent plots located in southwestern Amazonia, we calculated the mortality rate for three main modes of tree death: standing, broken and uprooted. We thus identified the predominant mode of death over a 14 year period (2002–2016). We found that trees in the southwestern Amazon died mainly standing (325 trees, 0.8% year−1) and broken (362 trees, 0.8% year−1); significantly fewer trees died uprooted (156 trees, 0.4% year−1, equivalent to less than one in five of all trees dying). During the study period, the tree mode of death with the greatest proportion in the region alternated between standing and broken trees. Forest characteristics of the southwestern Amazon, like presence and high density of bamboo culms, and the fact that the region was subject to severe droughts in 2005 and 2010, may be affecting how trees die in southwestern Amazon. The presence of these factors makes the forest dynamics of the southwestern Amazon different from other regions of the Amazon basin.

The Amazon forest represents one of the world’s largest terrestrial carbon reservoirs. Here, we evaluated the role of soil texture, climate, vegetation, and distance to savanna on the distribution and stocks of soil pyrogenic carbon (PyC) in intact forests with no history of recent fire spanning the southern Amazonia forest-Cerrado Zone of Transition (ZOT). In 19 one hectare forest plots, including three Amazonian Dark Earth (ADE, terra preta) sites with high soil PyC, we measured all trees and lianas with diameter ≥ 10 cm and analyzed soil physicochemical properties, including texture and PyC stocks. We quantified PyC stocks as a proportion of total organic carbon using hydrogen pyrolysis. We used multiple linear regression and variance partitioning to determine which variables best explain soil PyC variation. For all forests combined, soil PyC stocks ranged between 0.9 and 6.8 Mg/ha to 30 cm depth (mean 2.3 ± 1.5 Mg/ha) and PyC, on average, represented 4.3% of the total soil organic carbon (SOC). The most parsimonious model (based on AICc) included soil clay content and above-ground biomass (AGB) as the main predictors, explaining 71% of soil PyC variation. After removal of the ADE plots, PyC stocks ranged between 0.9 and 3.8 Mg/ha (mean 1.9 ± 0.8 Mg/ha–1) and PyC continued to represent ∼4% of the total SOC. The most parsimonious models without ADE included AGB and sand as the best predictors, with sand and PyC having an inverse relationship, and sand explaining 65% of the soil PyC variation. Partial regression analysis did not identify any of the components (climatic, environmental, and edaphic), pure or shared, as important in explaining soil PyC variation with or without ADE plots. We observed a substantial amount of soil PyC, even excluding ADE forests; however, contrary to expectations, soil PyC stocks were not higher nearer to the fire-dependent Cerrado than more humid regions of Amazonia. Our findings that soil texture and AGB explain the distribution and amount of soil PyC in ZOT forests will help to improve model estimates of SOC change with further climatic warming.

Data capturing multiple axes of tree size and shape, such as a tree's stem diameter, height and crown size, underpin a wide range of ecological research-from developing and testing theory on forest structure and dynamics, to estimating forest carbon stocks and their uncertainties, and integrating remote sensing imagery into forest monitoring programmes. However, these data can be surprisingly hard to come by, particularly for certain regions of the world and for specific taxonomic groups, posing a real barrier to progress in these fields. To overcome this challenge, we developed the Tallo database, a collection of 498,838 georeferenced and taxonomically standardized records of individual trees for which stem diameter, height and/or crown radius have been measured. These data were collected at 61,856 globally distributed sites, spanning all major forested and non-forested biomes. The majority of trees in the database are identified to species (88%), and collectively Tallo includes data for 5163 species distributed across 1453 genera and 187 plant families. The database is publicly archived under a CC-BY 4.0 licence and can be access from: https://doi.org/10.5281/zenodo.6637599. To demonstrate its value, here we present three case studies that highlight how the Tallo database can be used to address a range of theoretical and applied questions in ecology-from testing the predictions of metabolic scaling theory, to exploring the limits of tree allometric plasticity along environmental gradients and modelling global variation in maximum attainable tree height. In doing so, we provide a key resource for field ecologists, remote sensing researchers and the modelling community working together to better understand the role that trees play in regulating the terrestrial carbon cycle.

One of the most fundamental questions in ecology is how many species inhabit the Earth. However, due to massive logistical and financial challenges and taxonomic difficulties connected to the species concept definition, the global numbers of species, including those of important and well-studied life forms such as trees, still remain largely unknown. Here, based on global ground-sourced data, we estimate the total tree species richness at global, continental, and biome levels. Our results indicate that there are ∼73,000 tree species globally, among which ∼9,000 tree species are yet to be discovered. Roughly 40% of undiscovered tree species are in South America. Moreover, almost one-third of all tree species to be discovered may be rare, with very low populations and limited spatial distribution (likely in remote tropical lowlands and mountains). These findings highlight the vulnerability of global forest biodiversity to anthropogenic changes in land use and climate, which disproportionately threaten rare species and thus, global tree richness.

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

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.

Non-structural carbohydrates (NSC) are major substrates for plant metabolism and have been implicated in mediating drought-induced tree mortality. Despite their significance, NSC dynamics in tropical forests remain little studied. We present leaf and branch NSC data for 82 Amazon canopy tree species in six sites spanning a broad precipitation gradient. During the wet season, total NSC (NSCT) concentrations in both organs were remarkably similar across communities. However, NSCT and its soluble sugar (SS) and starch components varied much more across sites during the dry season. Notably, the proportion of leaf NSCT in the form of SS (SS:NSCT) increased greatly in the dry season in almost all species in the driest sites, implying an important role of SS in mediating water stress in these sites. This adjustment of leaf NSC balance was not observed in tree species less-adapted to water deficit, even under exceptionally dry conditions. Thus, leaf carbon metabolism may help to explain floristic sorting across water availability gradients in Amazonia and enable better prediction of forest responses to future climate change.

The responses of tropical forests to environmental change are critical uncertainties in predicting the future impacts of climate change. The positive phase of the 2015-2016 El Niño Southern Oscillation resulted in unprecedented heat and low precipitation in the tropics with substantial impacts on the global carbon cycle. The role of African tropical forests is uncertain as their responses to short-term drought and temperature anomalies have yet to be determined using on-the-ground measurements. African tropical forests may be particularly sensitive because they exist in relatively dry conditions compared with Amazonian or Asian forests, or they may be more resistant because of an abundance of drought-adapted species. Here, we report responses of structurally intact old-growth lowland tropical forests inventoried within the African Tropical Rainforest Observatory Network (AfriTRON). We use 100 long-term inventory plots from six countries each measured at least twice prior to and once following the 2015-2016 El Niño event. These plots experienced the highest temperatures and driest conditions on record. The record temperature did not significantly reduce carbon gains from tree growth or significantly increase carbon losses from tree mortality, but the record drought did significantly decrease net carbon uptake. Overall, the long-term biomass increase of these forests was reduced due to the El Niño event, but these plots remained a live biomass carbon sink (0.51 ± 0.40 Mg C ha-1 y-1) despite extreme environmental conditions. Our analyses, while limited to African tropical forests, suggest they may be more resistant to climatic extremes than Amazonian and Asian forests.

Tropical forests are the most diverse and productive ecosystems on Earth. While better understanding of these forests is critical for our collective future, until quite recently efforts to measure and monitor them have been largely disconnected. Networking is essential to discover the answers to questions that transcend borders and the horizons of funding agencies. Here we show how a global community is responding to the challenges of tropical ecosystem research with diverse teams measuring forests tree-by-tree in thousands of long-term plots. We review the major scientific discoveries of this work and show how this process is changing tropical forest science. Our core approach involves linking long-term grassroots initiatives with standardized protocols and data management to generate robust scaled-up results. By connecting tropical researchers and elevating their status, our Social Research Network model recognises the key role of the data originator in scientific discovery. Conceived in 1999 with RAINFOR (South America), our permanent plot networks have been adapted to Africa (AfriTRON) and Southeast Asia (T-FORCES) and widely emulated worldwide. Now these multiple initiatives are integrated via ForestPlots.net cyber-infrastructure, linking colleagues from 54 countries across 24 plot networks. Collectively these are transforming understanding of tropical forests and their biospheric role. Together we have discovered how, where and why forest carbon and biodiversity are responding to climate change, and how they feedback on it. This long-term pan-tropical collaboration has revealed a large long-term carbon sink and its trends, as well as making clear which drivers are most important, which forest processes are affected, where they are changing, what the lags are, and the likely future responses of tropical forests as the climate continues to change. By leveraging a remarkably old technology, plot networks are sparking a very modern revolution in tropical forest science. In the future, humanity can benefit greatly by nurturing the grassroots communities now collectively capable of generating unique, long-term understanding of Earth's most precious forests. Resumen: Los bosques tropicales son los ecosistemas más diversos y productivos del mundo y entender su funcionamiento es crítico para nuestro futuro colectivo. Sin embargo, hasta hace muy poco, los esfuerzos para medirlos y monitorearlos han estado muy desconectados. El trabajo en redes es esencial para descubrir las respuestas a preguntas que trascienden las fronteras y los plazos de las agencias de financiamiento. Aquí mostramos cómo una comunidad global está respondiendo a los desafíos de la investigación en ecosistemas tropicales a través de diversos equipos realizando mediciones árbol por árbol en miles de parcelas permanentes de largo plazo. Revisamos los descubrimientos más importantes de este trabajo y discutimos cómo este proceso está cambiando la ciencia relacionada a los bosques tropicales. El enfoque central de nuestro esfuerzo implica la conexión de iniciativas locales de largo plazo con protocolos estandarizados y manejo de datos para producir resultados que se puedan trasladar a múltiples escalas. Conectando investigadores tropicales, elevando su posición y estatus, nuestro modelo de Red Social de Investigación reconoce el rol fundamental que tienen, para el descubrimiento científico, quienes generan o producen los datos. Concebida en 1999 con RAINFOR (Suramérica), nuestras redes de parcelas permanentes han sido adaptadas en África (AfriTRON) y el sureste asiático (T-FORCES) y ampliamente replicadas en el mundo. Actualmente todas estas iniciativas están integradas a través de la ciber-infraestructura de ForestPlots.net, conectando colegas de 54 países en 24 redes diferentes de parcelas. Colectivamente, estas redes están transformando nuestro conocimiento sobre los bosques tropicales y el rol de éstos en la biósfera. Juntos hemos descubierto cómo, dónde y porqué el carbono y la biodiversidad de los bosques tropicales está respondiendo al cambio climático y cómo se retroalimentan. Esta colaboración pan-tropical de largo plazo ha expuesto un gran sumidero de carbono y sus tendencias, mostrando claramente cuáles son los factores más importantes, qué procesos se ven afectados, dónde ocurren los cambios, los tiempos de reacción y las probables respuestas futuras mientras el clima continúa cambiando. Apalancando lo que realmente es una tecnología antigua, las redes de parcelas están generando una verdadera y moderna revolución en la ciencia tropical. En el futuro, la humanidad puede beneficiarse enormemente si se nutren y cultivan comunidades de investigadores de base, actualmente con la capacidad de generar información única y de largo plazo para entender los que probablemente son los bosques más preciados de la tierra. Resumo: Florestas tropicais são os ecossistemas mais diversos e produtivos da Terra. Embora uma boa compreensão destas florestas seja crucial para o nosso futuro coletivo, até muito recentemente os esforços de medições e monitoramento tem sido amplamente desconexos. É essencial formarmos redes para obtermos respostas que transcendam as fronteiras e horizontes das agências financiadoras. Neste estudo nós mostramos como uma comunidade global está respondendo aos desafios da pesquisa de ecossistemas tropicais, com equipes diversas medindo florestas, árvore por árvore, em milhares de parcelas monitoradas a longo prazo. Nós revisamos as maiores descobertas científicas deste esforço global, e mostramos também como este processo vem mudando a ciência de florestas tropicais. Nossa abordagem principal envolve unir iniciativas de base a protocolos padronizados e gerenciamento de dados a fim de gerar resultados robustos em grandes escalas. Ao conectar pesquisadores tropicais e elevar seus status, nosso modelo de Rede de Pesquisa Social reconhece o papel chave do produtor dos dados na descoberta científica. Concebida em 1999 com o RAINFOR (América do Sul), nossa rede de parcelas permanentes foi adaptada para África (AfriTRON) e Sudeste Asiático (T-FORCES), e tem sido extensamente reproduzida em todo o mundo. Agora estas múltiplas iniciativas estão integradas através da infraestrutura cibernética do ForestPlots.net, conectando colegas de 54 países e 24 redes de parcelas. Estas iniciativas estão transformando coletivamente o entendimento das florestas tropicais e seus papéis na biosfera. Juntos nós descobrimos como, onde e por que o carbono e a biodiversidade da floresta estão respondendo às mudanças climáticas, e seus efeitos de retroalimentação. Esta duradoura colaboração pantropical revelou um grande sumidouro de carbono persistente e suas tendências, assim como tem evidenciado quais os fatores que influenciam essas tendências, quais processos florestais são mais afetados, onde eles estão mudando, seus atrasos no tempo de resposta, e as prováveis respostas das florestas tropicais conforme o clima continua a mudar. Dessa forma, aproveitando uma notável tecnologia antiga, redes de parcelas acendem as faíscas de uma moderna revolução na ciência das florestas tropicais. No futuro a humanidade pode se beneficiar incentivando estas comunidades locais que agora são coletivamente capazes de gerar conhecimentos únicos e duradouros sobre as florestas mais preciosas da Terra. Résume: Les forêts tropicales sont les écosystèmes les plus diversifiés et les plus productifs de la planète. Si une meilleure compréhension de ces forêts est essentielle pour notre avenir collectif, jusqu'à tout récemment, les efforts déployés pour les mesurer et les surveiller ont été largement déconnectés. La mise en réseau est essentielle pour découvrir les réponses à des questions qui dépassent les frontières et les horizons des organismes de financement. Nous montrons ici comment une communauté mondiale relève les défis de la recherche sur les écosystèmes tropicaux avec diverses équipes qui mesurent les forêts arbre après arbre dans de milliers de parcelles permanentes. Nous passons en revue les principales découvertes scientifiques de ces travaux et montrons comment ce processus modifie la science des forêts tropicales. Notre approche principale consiste à relier les initiatives de base à long terme à des protocoles standardisés et une gestion de données afin de générer des résultats solides à grande échelle. En reliant les chercheurs tropicaux et en élevant leur statut, notre modèle de réseau de recherche sociale reconnaît le rôle clé de l'auteur des données dans la découverte scientifique. Conçus en 1999 avec RAINFOR (Amérique du Sud), nos réseaux de parcelles permanentes ont été adaptés à l'Afrique (AfriTRON) et à l'Asie du Sud-Est (T-FORCES) et largement imités dans le monde entier. Ces multiples initiatives sont désormais intégrées via l'infrastructure ForestPlots.net, qui relie des collègues de 54 pays à travers 24 réseaux de parcelles. Ensemble, elles transforment la compréhension des forêts tropicales et de leur rôle biosphérique. Ensemble, nous avons découvert comment, où et pourquoi le carbone forestier et la biodiversité réagissent au changement climatique, et comment ils y réagissent. Cette collaboration pan-tropicale à long terme a révélé un important puits de carbone à long terme et ses tendances, tout en mettant en évidence les facteurs les plus importants, les processus forestiers qui sont affectés, les endroits où ils changent, les décalages et les réactions futures probables des forêts tropicales à mesure que le climat continue de changer. En tirant parti d'une technologie remarquablement ancienne, les réseaux de parcelles déclenchent une révolution très moderne dans la science des forêts tropicales. À l'avenir, l'humanité pourra grandement bénéficier du soutien des communautés de base qui sont maintenant collectivement capables de générer une compréhension unique et à long terme des forêts les plus précieuses de la Terre. Abstrak: Hutan tropika adalah di antara ekosistem yang paling produktif dan mempunyai kepelbagaian biodiversiti yang tinggi di seluruh dunia. Walaupun pemahaman mengenai hutan tropika amat penting untuk masa depan kita, usaha-usaha untuk mengkaji dan mengawas hutah-hutan tersebut baru sekarang menjadi lebih diperhubungkan. Perangkaian adalah sangat penting untuk mencari jawapan kepada soalan-soalan yang menjangkaui sempadan dan batasan agensi pendanaan. Di sini kami menunjukkan bagaimana sebuah komuniti global bertindak balas terhadap cabaran penyelidikan ekosistem tropika melalui penglibatan pelbagai kumpulan yang mengukur hutan secara pokok demi pokok dalam beribu-ribu plot jangka panjang. Kami meninjau semula penemuan saintifik utama daripada kerja ini dan menunjukkan bagaimana proses ini sedang mengubah bidang sains hutan tropika. Teras pendekatan kami memberi tumpuan terhadap penghubungan inisiatif akar umbi jangka panjang dengan protokol standar serta pengurusan data untuk mendapatkan hasil skala besar yang kukuh. Dengan menghubungkan penyelidik-penyelidik tropika dan meningkatkan status mereka, model Rangkaian Penyelidikan Sosial kami mengiktiraf kepentingan peranan pengasas data dalam penemuan saintifik. Bermula dengan pengasasan RAINFOR (Amerika Selatan) pada tahun 1999, rangkaian-rangkaian plot kekal kami kemudian disesuaikan untuk Afrika (AfriTRON) dan Asia Tenggara (T-FORCES) dan selanjutnya telah banyak dicontohi di seluruh dunia. Kini, inisiatif-inisiatif tersebut disepadukan melalui infrastruktur siber ForestPlots.net yang menghubungkan rakan sekerja dari 54 negara di 24 buah rangkaian plot. Secara kolektif, rangkaian ini sedang mengubah pemahaman tentang hutan tropika dan peranannya dalam biosfera. Kami telah bekerjasama untuk menemukan bagaimana, di mana dan mengapa karbon serta biodiversiti hutan bertindak balas terhadap perubahan iklim dan juga bagaimana mereka saling bermaklum balas. Kolaborasi pan-tropika jangka panjang ini telah mendedahkan sebuah sinki karbon jangka panjang serta arah alirannya dan juga menjelaskan pemandu-pemandu perubahan yang terpenting, di mana dan bagaimana proses hutan terjejas, masa susul yang ada dan kemungkinan tindakbalas hutan tropika pada perubahan iklim secara berterusan di masa depan. Dengan memanfaatkan pendekatan lama, rangkaian plot sedang menyalakan revolusi yang amat moden dalam sains hutan tropika. Pada masa akan datang, manusia sejagat akan banyak mendapat manfaat jika memupuk komuniti-komuniti akar umbi yang kini berkemampuan secara kolektif menghasilkan pemahaman unik dan jangka panjang mengenai hutan-hutan yang paling berharga di dunia.

Aims: We conducted a suite of tracer studies using the stable isotope 13C to follow and quantify the flow of carbon from leaf litter and roots into soil components including aggregates and biota with and without invasive earthworms. Methods: Ten-year-old saplings of sugar maple growing in the understory of a thinned northern hardwood forest were labeled with 13CO2. The 13C labeled leaf litter was applied to forest plots with and without invasive earthworms (Lumbricidae) and traced for three years. We also traced the label from the trees through the roots and into soil components in the labeling chambers. Labeled fine roots and stem wood were incubated in a forest and the label was quantified over six years of decomposition. Results: We were able to detect the litter tracer to 10 cm soil depth in plots without earthworms and to 20 cm with earthworms present, and earthworms promoted C incorporation into soil aggregates. The soil food web was much more enriched in the label from roots than from aboveground plant litter. Rapid fine root decay was observed (k = 0.9 yr−1), and although labelled wood was almost completely decayed, little 13C was recovered in soil (0.33%). Conclusion: the approach was successful for quantifying transport and fate of tree carbon in forest soils and could be enhanced with careful quantification of gross assimilation.

Botanical assessment was conducted in the Boabeng-Fiema Monkey Sanctuary (BFMS), aimed at assessing the woody plant composition, diversity and structure, of vegetation types and conservation importance of the reserve. Ten (20m×50m) plots were established in forest, transition and savanna stands of BFMS. Overall, 66 species representing 55 genera and 26 families were recorded. Floristic attributes showed lower number of species, families, genera and stem density in savanna (24, 14, 21 and 97, respectively) compared to transition (37, 21, 35, 596) and forest (42, 20, 37, 149). The most dominant families were Fabaceae, Moraceae, Rubiaceae and Malvaceae whilst trees were the most predominant growth form accounting for 66% of species recorded. Trees were more diverse in forest (1.32) and savanna (1.10) while shrubs were more diverse in transition (1.14). Non-metric multidimensional scaling ordination showed distribution of distinct tree and shrub species across stands. Anogeisus leiocarpus and Detarium senegalensis were predominant in the savanna vegetation with Importance Value Indices (IVI) of 93.1 and 34.9, respectively, while in the transition zone, A. leiocarpus and the shrub, Uvaria chamae, were the most dominant with IVI of 71.1 and 38.9, respectively. Dominant species in the forest were Bombax buonopozense and Lecaniodiscus cupanioides with IVI of 42.5 and 25.1, respectively. The transition zone had a Genetic Heat Index (GHI) of 254.3, followed by the forest and savanna zones with values of 140.0 and 97.6, respectively. Our results indicated the BFMS harbours several valuable plant species, suggesting its potential as in-situ conservation as well as tourist, research and education site.

Amazonian forests are extraordinarily diverse, but the estimated species richness is very much debated. Here, we apply an ensemble of parametric estimators and a novel technique that includes conspecific spatial aggregation to an extended database of forest plots with up-to-date taxonomy. We show that the species abundance distribution of Amazonia is best approximated by a logseries with aggregated individuals, where aggregation increases with rarity. By averaging several methods to estimate total richness, we confirm that over 15,000 tree species are expected to occur in Amazonia. We also show that using ten times the number of plots would result in an increase to just ~50% of those 15,000 estimated species. To get a more complete sample of all tree species, rigorous field campaigns may be needed but the number of trees in Amazonia will remain an estimate for years to come.

Lianas, a key component of tropical forests, can limit growth of trees, impacting both the structure and functioning of forests, and are expected to benefit from fragmentation and potentially from global climatic changes. While it is critical to understand the impacts of liana infestation on contemporary tropical forests across large geographical areas, to date most liana studies have been focussed on single or few sites. We measured and quantified liana infestation of 16,066 trees with diameter ≥10 cm in 27 plots distributed across southern Amazonia, a region characterized by substantial ecological and environmental variation and environmental change. We examined the influence of potential drivers of liana infestation at the plot, species and individual tree level. Additionally, we evaluated the effect of liana infestation on tree growth. More than half of all trees had lianas in their crown. At the plot level, infestation by lianas was driven by forest structure but not by the studied climate or soil fertility variables, though low levels of liana infestation were found in seasonally flooded forests. At the tree level, larger and stouter trees had a greater proportion of their crown infested with lianas. At the species level, trees belonging to intrinsically slow-growing, dense-wooded species were more susceptible to liana infestation. Liana infestation had a negative impact on tree growth, with growth of heavily infested trees reduced by 33% compared to non-infested. The impact of liana infestation on tree growth was strongest for the best-lit tree crowns, indicating that lianas act to reduce the large competitive advantage that well-lit trees otherwise hold over their neighbours. Synthesis. Lianas are a pervasive and influential feature of the extensive forests at the southern edge of Amazonia. The degree of liana infestation in forests was closely linked to species-level variables such as potential growth and wood density as well as the size of the individual tree. The growth of heavily infested trees was particularly restricted by lianas, and especially so for trees growing in otherwise favourable conditions, indicating the potential for lianas not only to reduce forest growth rates overall, but also to modify competitive hierarchies among trees within tropical forests.

Competition among trees is an important driver of community structure and dynamics in tropical forests. Neighboring trees may impact an individual tree's growth rate and probability of mortality, but large-scale geographic and environmental variation in these competitive effects has yet to be evaluated across the tropical forest biome. We quantified effects of competition on tree-level basal area growth and mortality for trees ≥10-cm diameter across 151 ~1-ha plots in mature tropical forests in Amazonia and tropical Africa by developing nonlinear models that accounted for wood density, tree size, and neighborhood crowding. Using these models, we assessed how water availability (i.e. climatic water deficit) and soil fertility influenced the predicted plot-level strength of competition (i.e. the extent to which growth is reduced, or mortality is increased, by competition across all individual trees). On both continents, tree basal area growth decreased with wood density and increased with tree size. Growth decreased with neighborhood crowding, which suggests that competition is important. Tree mortality decreased with wood density and generally increased with tree size, but was apparently unaffected by neighborhood crowding. Across plots, variation in the plot-level strength of competition was most strongly related to plot basal area (i.e. the sum of the basal area of all trees in a plot), with greater reductions in growth occurring in forests with high basal area, but in Amazonia, the strength of competition also varied with plot-level wood density. In Amazonia, the strength of competition increased with water availability because of the greater basal area of wetter forests, but was only weakly related to soil fertility. In Africa, competition was weakly related to soil fertility and invariant across the shorter water availability gradient. Overall, our results suggest that competition influences the structure and dynamics of tropical forests primarily through effects on individual tree growth rather than mortality and that the strength of competition largely depends on environment-mediated variation in basal area.

The long-term dynamics of regeneration in tropical forests dominated by single tree species remains largely undocumented, yet is key to understanding the mechanisms by which one species can gain dominance and resist environmental change. We report here on the long-term regeneration dynamics in a monodominant stand of Brosimum rubescens Taub. (Moraceae) at the southern border of the Amazon forest. Here the climate has warmed and dried since the mid-1990′s. Twenty-one years of tree and liana regeneration were evaluated in four censuses in 30 plots by assessing species abundance, dominance, and diversity in all regeneration classes up to 5 cm diameter. The density of B. rubescens seedlings declined markedly, from 85% in 1997 to 29% in 2018 after the most intense El Niño-driven drought. While the fraction contributed by other tree species changed little, the relative density of liana seedlings increased from just 1 to 54% and three-quarters of liana species underwent a ten-fold or greater increase in abundance. The regeneration community experienced a high rate of species turnover, with changes in the overall richness and species diversity determined principally by lianas, not trees. Long-term maintenance of monodominance in this tropical forest is threatened by a sharp decline in the regeneration of the monodominant species and the increase in liana density, suggesting that monodominance will prove to be a transitory condition. The close association of these rapid changes with drying indicates that monodominant B. rubescens forests are impacted by drought-driven changes in regeneration, and therefore are particularly sensitive to climatic change.

El Niño years are characterized by a high sea surface temperature anomaly in the Equatorial Pacific Ocean, which leads to unusually warm and dry conditions over many fire-prone regions globally. This can lead to an increase in burned area and emissions from fire activity, and socio-economic, and environmental losses. Previous studies using satellite observations to assess the impacts of the recent 2015/16 El Niño found an increase in burned area in some regions compared to La Niña years. Here, we use the dynamic land surface model JULES to assess how conditions differed as a result of the El Niño by comparing simulations driven by observations from the year 2015/16 with mean climatological drivers of temperature, precipitation, humidity, wind, air pressure, and short and long-wave radiation. We use JULES with the interactive fire module INFERNO to assess the effects on precipitation, temperature, burned area, and the associated impacts on the carbon sink globally and for three regions: South America, Africa, and Asia. We find that the model projects a variable response in precipitation, with some areas including northern South America, southern Africa and East Asia getting drier, and most areas globally seeing an increase in temperature. As a result, higher burned area is simulated with El Niño conditions in most regions, although there are areas of both increased and decreased burned area over Africa. South America shows the largest fire response with El Niño, with a 13% increase in burned area and emitted carbon, corresponding with the largest decrease in carbon uptake. Within South America, peak fire occurs from August to October across central-southern Brazil, and temperature is shown to be the main driver of the El Niño-induced increase in burned area during this period. Combined, our results indicate that although 2015/16 was not a peak year for global total burned area or fire emissions, the El Niño led to an overall increase of 4% in burned area and 5% in emissions compared to a “No El Niño” scenario for 2015/16, and contributed to a 4% reduction in the terrestrial carbon sink.

Fire in tropical forests increases tree mortality, degrades forest structure, and reduces carbon stocks. Currently, there are large gaps in understanding how fire affects understory forest structure and composition, interactions with fire recurrence, and long-term impacts. Understanding these changes is critical to evaluate the present and future response of tropical forests to fire. We studied post-fire changes in understory regeneration in forests in Mato Grosso State, southern Amazonia, Brazil, aiming to answer the following questions: (i) does forest structure (basal area) and tree community composition vary with fire frequency and time since the last fire? (ii) does the response differ among strata (e.g. sapling, larger trees)? (iii) are changes in diversity associated with changes in forest structure? We surveyed trees and lianas in previously structurally intact forests that underwent selective logging, followed by different fire histories, including 5 and 16 years after once-burned, 5 years after three times burned, and unburned (control). Overall, species composition (abundance, richness, and number of families) and diversity were highest for the unburned treatment and lowest for the recurrent burned areas. Fire frequency negatively affected plant structure and basal area; basal area of small, medium, and large plants declined significantly by more than 50% in the most frequently burned areas. Richness was positively related to basal area in the three times burned sites and in the 16 years regenerating site for all strata. Our results demonstrate the negative influence of frequent fires on both the composition and structure of small trees in Amazonian forest. These changes to the cohort of small-sized trees may persist and have long-term impacts on forest structure, affecting the capacity, and direction of forest recovery. With wildfire widespread across the region and increasing in frequency, fire may negatively affect tree diversity in remaining selectively logged forests, and affect regional carbon cycling with consequences for the global vegetation carbon sink.

The sensitivity of tropical forest carbon to climate is a key uncertainty in predicting global climate change. Although short-term drying and warming are known to affect forests, it is unknown if such effects translate into long-term responses. Here, we analyze 590 permanent plots measured across the tropics to derive the equilibrium climate controls on forest carbon. Maximum temperature is the most important predictor of aboveground biomass (-9.1 megagrams of carbon per hectare per degree Celsius), primarily by reducing woody productivity, and has a greater impact per °C in the hottest forests (>32.2°C). Our results nevertheless reveal greater thermal resilience than observations of short-term variation imply. To realize the long-term climate adaptation potential of tropical forests requires both protecting them and stabilizing Earth's climate.

Background and aims: We explored the hypothesis that low soil water-holding capacity is the main factor driving the monodominance of Brosimum rubescens in a monodominant forest in Southern Amazonia. Tropical monodominant forests are rare ecosystems with low diversity and high dominance of a single tree species. The causes of this atypical condition are still poorly understood. Some studies have shown a relationship between monodominance and waterlogging or soil attributes, while others have concluded that edaphic factors have little or no explanatory value, but none has accounted for soil-moisture variation other than waterlogging. This study is the first to explicitly explore how low soil water-holding capacity influences the monodominance of tropical forests. Methods: We conducted in situ measurements of vertical soil moisture using electrical resistance collected over 1 year at 0–5; 35–40 and 75–80 cm depths in a B. rubescens monodominant forest and in an adjacent mixed-species forest in the Amazon-Cerrado transition zone, Brazil. Minimum leaf water potential (Ψmin) of the seven most common species, including B. rubescens, and soil water-holding capacity for both forests were determined. Results: the vertical soil moisture decay pattern was similar in both forests for all depths. However, the slightly higher water availability in the monodominant forest and Ψmin similarity between B. rubescens and nearby mixed forest species indicate that low water-availability does not cause the monodominance. Conclusions: We reject the hypothesis that monodominance of B. rubescens is primarily determined by low soil water-holding capacity, reinforcing the idea that monodominance in tropical forests is not determined by a single factor.

Aim: Palms are an iconic, diverse and often abundant component of tropical ecosystems that provide many ecosystem services. Being monocots, tree palms are evolutionarily, morphologically and physiologically distinct from other trees, and these differences have important consequences for ecosystem services (e.g. carbon sequestration and storage) and in terms of responses to climate change. We quantified global patterns of tree palm relative abundance to help improve understanding of tropical forests and reduce uncertainty about these ecosystems under climate change. Location: Tropical and subtropical moist forests. Time period: Current. Major taxa studied: Palms (Arecaceae). Methods: We assembled a pantropical dataset of 2,548 forest plots (covering 1,191 ha) and quantified tree palm (i.e. ≥10 cm diameter at breast height) abundance relative to co-occurring non-palm trees. We compared the relative abundance of tree palms across biogeographical realms and tested for associations with palaeoclimate stability, current climate, edaphic conditions and metrics of forest structure. Results: on average, the relative abundance of tree palms was more than five times larger between Neotropical locations and other biogeographical realms. Tree palms were absent in most locations outside the Neotropics but present in >80% of Neotropical locations. The relative abundance of tree palms was more strongly associated with local conditions (e.g. higher mean annual precipitation, lower soil fertility, shallower water table and lower plot mean wood density) than metrics of long-term climate stability. Life-form diversity also influenced the patterns; palm assemblages outside the Neotropics comprise many non-tree (e.g. climbing) palms. Finally, we show that tree palms can influence estimates of above-ground biomass, but the magnitude and direction of the effect require additional work. Conclusions: Tree palms are not only quintessentially tropical, but they are also overwhelmingly Neotropical. Future work to understand the contributions of tree palms to biomass estimates and carbon cycling will be particularly crucial in Neotropical forests.

Less than half of the original two million square kilometers of the Cerrado vegetation remains standing, and there are still many uncertainties as to how to conserve and prioritize remaining areas effectively. A key limitation is the continuing lack of geographically-extensive evaluation of ecosystem-level properties across the biome. Here we sought to address this gap by comparing the woody vegetation of the typical cerrado of the Cerrado–Amazonia Transition with that of the core area of the Cerrado in terms of both tree diversity and vegetation biomass. We used 21 one-hectare plots in the transition and 18 in the core to compare key structural parameters (tree height, basal area, and above-ground biomass), and diversity metrics between the regions. We also evaluated the effects of temperature and precipitation on biomass, as well as explored the species diversity versus biomass relationship. We found, for the first time, both that the typical cerrado at the transition holds substantially more biomass than at the core, and that higher temperature and greater precipitation can explain this difference. By contrast, plot-level alpha diversity was almost identical in the two regions. Finally, contrary to some theoretical expectations, we found no positive relationship between species diversity and biomass for the Cerrado woody vegetation. This has implications for the development of effective conservation measures, given that areas with high biomass and importance for the compensation of greenhouse gas emissions are often not those with the greatest diversity.

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 identity of the dominant root-associated microbial symbionts in a forest determines the ability of trees to access limiting nutrients from atmospheric or soil pools1,2, sequester carbon3,4 and withstand the effects of climate change5,6. Characterizing the global distribution of these symbioses and identifying the factors that control this distribution are thus integral to understanding the present and future functioning of forest ecosystems. Here we generate a spatially explicit global map of the symbiotic status of forests, using a database of over 1.1 million forest inventory plots that collectively contain over 28,000 tree species. Our analyses indicate that climate variables—in particular, climatically controlled variation in the rate of decomposition—are the primary drivers of the global distribution of major symbioses. We estimate that ectomycorrhizal trees, which represent only 2% of all plant species7, constitute approximately 60% of tree stems on Earth. Ectomycorrhizal symbiosis dominates forests in which seasonally cold and dry climates inhibit decomposition, and is the predominant form of symbiosis at high latitudes and elevation. By contrast, arbuscular mycorrhizal trees dominate in aseasonal, warm tropical forests, and occur with ectomycorrhizal trees in temperate biomes in which seasonally warm-and-wet climates enhance decomposition. Continental transitions between forests dominated by ectomycorrhizal or arbuscular mycorrhizal trees occur relatively abruptly along climate-driven decomposition gradients; these transitions are probably caused by positive feedback effects between plants and microorganisms. Symbiotic nitrogen fixers—which are insensitive to climatic controls on decomposition (compared with mycorrhizal fungi)—are most abundant in arid biomes with alkaline soils and high maximum temperatures. The climatically driven global symbiosis gradient that we document provides a spatially explicit quantitative understanding of microbial symbioses at the global scale, and demonstrates the critical role of microbial mutualisms in shaping the distribution of plant species.

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.

Climatic changes have profound effects on the distribution of biodiversity, but untangling the links between climatic change and ecosystem functioning is challenging, particularly in high diversity systems such as tropical forests. Tropical forests may also show different responses to a changing climate, with baseline climatic conditions potentially inducing differences in the strength and timing of responses to droughts. Trait-based approaches provide an opportunity to link functional composition, ecosystem function and environmental changes. We demonstrate the power of such approaches by presenting a novel analysis of long-term responses of different tropical forest to climatic changes along a rainfall gradient. We explore how key ecosystem's biogeochemical properties have shifted over time as a consequence of multi-decadal drying. Notably, we find that drier tropical forests have increased their deciduous species abundance and generally changed more functionally than forests growing in wetter conditions, suggesting an enhanced ability to adapt ecologically to a drying environment.

As countries advance in greenhouse gas (GHG) accounting for climate change mitigation, consistent estimates of aboveground net biomass change (∆AGB) are needed. Countries with limited forest monitoring capabilities in the tropics and subtropics rely on IPCC 2006 default ∆AGB rates, which are values per ecological zone, per continent. Similarly, research into forest biomass change at a large scale also makes use of these rates. IPCC 2006 default rates come from a handful of studies, provide no uncertainty indications and do not distinguish between older secondary forests and old-growth forests. As part of the 2019 Refinement to the 2006 IPCC Guidelines for National Greenhouse Gas Inventories, we incorporate ∆AGB data available from 2006 onwards, comprising 176 chronosequences in secondary forests and 536 permanent plots in old-growth and managed/logged forests located in 42 countries in Africa, North and South America and Asia. We generated ∆AGB rate estimates for younger secondary forests (≤20 years), older secondary forests (>20 years and up to 100 years) and old-growth forests, and accounted for uncertainties in our estimates. In tropical rainforests, for which data availability was the highest, our ∆AGB rate estimates ranged from 3.4 (Asia) to 7.6 (Africa) Mg ha-1  year-1 in younger secondary forests, from 2.3 (North and South America) to 3.5 (Africa) Mg ha-1  year-1 in older secondary forests, and 0.7 (Asia) to 1.3 (Africa) Mg ha-1  year-1 in old-growth forests. We provide a rigorous and traceable refinement of the IPCC 2006 default rates in tropical and subtropical ecological zones, and identify which areas require more research on ∆AGB. In this respect, this study should be considered as an important step towards quantifying the role of tropical and subtropical forests as carbon sinks with higher accuracy; our new rates can be used for large-scale GHG accounting by governmental bodies, nongovernmental organizations and in scientific research.

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.

Over recent decades, biomass gains in remaining old-growth Amazonia forests have declined due to environmental change. Amazonia's huge size and complexity makes understanding these changes, drivers, and consequences very challenging. Here, using a network of permanent monitoring plots at the Amazon-Cerrado transition, we quantify recent biomass carbon changes and explore their environmental drivers. Our study area covers 30 plots of upland and riparian forests sampled at least twice between 1996 and 2016 and subject to various levels of fire and drought. Using these plots, we aimed to: (1) estimate the long-term biomass change rate; (2) determine the extent to which forest changes are influenced by forest type; and (3) assess the threat to forests from ongoing environmental change. Overall, there was no net change in biomass, but there was clear variation among different forest types. Burning occurred at least once in 8 of the 12 riparian forests, while only 1 of the 18 upland forests burned, resulting in losses of carbon in burned riparian forests. Net biomass gains prevailed among other riparian and upland forests throughout Amazonia. Our results reveal an unanticipated vulnerability of riparian forests to fire, likely aggravated by drought, and threatening ecosystem conservation at the Amazon southern margins.

Climate, species composition, and soils are thought to control carbon cycling and forest structure in Amazonian forests. Here, we add a demographics scheme (tree recruitment, growth, and mortality) to a recently developed non-demographic model—the Trait-based Forest Simulator (TFS)—to explore the roles of climate and plant traits in controlling forest productivity and structure. We compared two sites with differing climates (seasonal vs. aseasonal precipitation) and plant traits. Through an initial validation simulation, we assessed whether the model converges on observed forest properties (productivity, demographic and structural variables) using datasets of functional traits, structure, and climate to model the carbon cycle at the two sites. In a second set of simulations, we tested the relative importance of climate and plant traits for forest properties within the TFS framework using the climate from the two sites with hypothetical trait distributions representing two axes of functional variation (“fast” vs. “slow” leaf traits, and high vs. low wood density). The adapted model with demographics reproduced observed variation in gross (GPP) and net (NPP) primary production, and respiration. However, NPP and respiration at the level of plant organs (leaf, stem, and root) were poorly simulated. Mortality and recruitment rates were underestimated. The equilibrium forest structure differed from observations of stem numbers suggesting either that the forests are not currently at equilibrium or that mechanisms are missing from the model. Findings from the second set of simulations demonstrated that differences in productivity were driven by climate, rather than plant traits. Contrary to expectation, varying leaf traits had no influence on GPP. Drivers of simulated forest structure were complex, with a key role for wood density mediated by its link to tree mortality. Modeled mortality and recruitment rates were linked to plant traits alone, drought-related mortality was not accounted for. In future, model development should focus on improving allocation, mortality, organ respiration, simulation of understory trees and adding hydraulic traits. This type of model that incorporates diverse tree strategies, detailed forest structure and realistic physiology is necessary if we are to be able to simulate tropical forest responses to global change scenarios.

Tropical forests are known for their high diversity. Yet, forest patches do occur in the tropics where a single tree species is dominant. Such “monodominant” forests are known from all of the main tropical regions. For Amazonia, we sampled the occurrence of monodominance in a massive, basin-wide database of forest-inventory plots from the Amazon Tree Diversity Network (ATDN). Utilizing a simple defining metric of at least half of the trees ≥ 10 cm diameter belonging to one species, we found only a few occurrences of monodominance in Amazonia, and the phenomenon was not significantly linked to previously hypothesized life history traits such wood density, seed mass, ectomycorrhizal associations, or Rhizobium nodulation. In our analysis, coppicing (the formation of sprouts at the base of the tree or on roots) was the only trait significantly linked to monodominance. While at specific locales coppicing or ectomycorrhizal associations may confer a considerable advantage to a tree species and lead to its monodominance, very few species have these traits. Mining of the ATDN dataset suggests that monodominance is quite rare in Amazonia, and may be linked primarily to edaphic factors.

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.

© 2019, the Author(s), under exclusive licence to Springer Nature Limited. Quantifying carbon dynamics in forests is critical for understanding their role in long-term climate regulation 1–4. Yet little is known about tree longevity in tropical forests 3,5–8 , a factor that is vital for estimating carbon persistence 3,4. Here we calculate mean carbon age (the period that carbon is fixed in trees 7 ) in different strata of African tropical forests using (1) growth-ring records with a unique timestamp accurately demarcating 66 years of growth in one site and (2) measurements of diameter increments from the African Tropical Rainforest Observation Network (23 sites). We find that in spite of their much smaller size, in understory trees mean carbon age (74 years) is greater than in sub-canopy (54 years) and canopy (57 years) trees and similar to carbon age in emergent trees (66 years). The remarkable carbon longevity in the understory results from slow and aperiodic growth as an adaptation to limited resource availability 9–11. Our analysis also reveals that while the understory represents a small share (11%) of the carbon stock 12,13 , it contributes disproportionally to the forest carbon sink (20%). We conclude that accounting for the diversity of carbon age and carbon sequestration among different forest strata is critical for effective conservation management 14–16 and for accurate modelling of carbon cycling 4.

Background: the remaining forests in the extensive contact zone between southern Amazonia (seasonal rain forest) and the Cerrado (savanna) biomes are at risk due to intense land-use and climate change. Aims: to explore the vulnerability of these transitional forests to changes in land use and climate, we evaluated the effects of fragmentation and climatic variables on forest structure. Methods: We measured the diameter and height of 14,185 trees with diameter ≥10 cm at 24 forest plots distributed over an area of 25,000 km 2. For each plot, we obtained data on contemporary fragmentation and climatic variables. Results: Forest structure variables (height, diameter, height:diameter allometry, biomass) varied significantly both within and among plots. The height, H:D and biomass of trees were positively correlated with annual precipitation and fragment area. Conclusions: the association between forest structure and precipitation indicates that these forests plots are likely to be vulnerable to dry season intensification anticipated for the southern edge of the Amazon. Additionally, the reduction in the fragment area may contribute to reductions in forest biomass and tree height, and consequently ecosystem carbon stocks. Given the likely susceptibility of these forests, urgent conservation action is needed to prevent further habitat degradation.

Quantifying the relationship between tree diameter and height is a key component of efforts to estimate biomass and carbon stocks in tropical forests. Although substantial site-to-site variation in height–diameter allometries has been documented, the time consuming nature of measuring all tree heights in an inventory plot means that most studies do not include height, or else use generic pan-tropical or regional allometric equations to estimate height. Using a pan-tropical dataset of 73 plots where at least 150 trees had in-field ground-based height measurements, we examined how the number of trees sampled affects the performance of locally derived height–diameter allometries, and evaluated the performance of different methods for sampling trees for height measurement. Using cross-validation, we found that allometries constructed with just 20 locally measured values could often predict tree height with lower error than regional or climate-based allometries (mean reduction in prediction error = 0.46 m). The predictive performance of locally derived allometries improved with sample size, but with diminishing returns in performance gains when more than 40 trees were sampled. Estimates of stand-level biomass produced using local allometries to estimate tree height show no over- or under-estimation bias when compared with biomass estimates using field measured heights. We evaluated five strategies to sample trees for height measurement, and found that sampling strategies that included measuring the heights of the ten largest diameter trees in a plot outperformed (in terms of resulting in local height–diameter models with low height prediction error) entirely random or diameter size-class stratified approaches. Our results indicate that even limited sampling of heights can be used to refine height–diameter allometries. We recommend aiming for a conservative threshold of sampling 50 trees per location for height measurement, and including the ten trees with the largest diameter in this sample.

Plant functional traits directly affect ecosystem functions. At the species level, trait combinations depend on trade-offs representing different ecological strategies, but at the community level trait combinations are expected to be decoupled from these trade-offs because different strategies can facilitate co-existence within communities. A key question is to what extent community-level trait composition is globally filtered and how well it is related to global versus local environmental drivers. Here, we perform a global, plot-level analysis of trait-environment relationships, using a database with more than 1.1 million vegetation plots and 26,632 plant species with trait information. Although we found a strong filtering of 17 functional traits, similar climate and soil conditions support communities differing greatly in mean trait values. The two main community trait axes that capture half of the global trait variation (plant stature and resource acquisitiveness) reflect the trade-offs at the species level but are weakly associated with climate and soil conditions at the global scale. Similarly, within-plot trait variation does not vary systematically with macro-environment. Our results indicate that, at fine spatial grain, macro-environmental drivers are much less important for functional trait composition than has been assumed from floristic analyses restricted to co-occurrence in large grid cells. Instead, trait combinations seem to be predominantly filtered by local-scale factors such as disturbance, fine-scale soil conditions, niche partitioning and biotic interactions.

Aim: Large tropical trees form the interface between ground and airborne observations, offering a unique opportunity to capture forest properties remotely and to investigate their variations on broad scales. However, despite rapid development of metrics to characterize the forest canopy from remotely sensed data, a gap remains between aerial and field inventories. To close this gap, we propose a new pan-tropical model to predict plot-level forest structure properties and biomass from only the largest trees. Location: Pan-tropical. Time period: Early 21st century. Major taxa studied: Woody plants. Methods: Using a dataset of 867 plots distributed among 118 sites across the tropics, we tested the prediction of the quadratic mean diameter, basal area, Lorey's height, community wood density and aboveground biomass (AGB) from the ith largest trees. Results: Measuring the largest trees in tropical forests enables unbiased predictions of plot- and site-level forest structure. The 20 largest trees per hectare predicted quadratic mean diameter, basal area, Lorey's height, community wood density and AGB with 12, 16, 4, 4 and 17.7% of relative error, respectively. Most of the remaining error in biomass prediction is driven by differences in the proportion of total biomass held in medium-sized trees (50–70 cm diameter at breast height), which shows some continental dependency, with American tropical forests presenting the highest proportion of total biomass in these intermediate-diameter classes relative to other continents. Main conclusions: Our approach provides new information on tropical forest structure and can be used to generate accurate field estimates of tropical forest carbon stocks to support the calibration and validation of current and forthcoming space missions. It will reduce the cost of field inventories and contribute to scientific understanding of tropical forest ecosystems and response to climate change.

In the “Cerrado”–Amazon ecotone in central Brazil, recent studies suggest some encroachment of forest into savanna, but how, where, and why this might be occurring is unclear. To better understand this phenomenon, we assessed changes in the structure and dynamics of tree species in three vegetation types at the “Cerrado”–Amazon ecotone that are potentially susceptible to encroachment: open “cerrado” (OC), typical “cerrado” (TC) and dense woodland (DW). We estimated changes in density, basal area and aboveground biomass of trees with diameter ≥ 10 cm over four inventories carried out between 2008 and 2015 and classified the species according to their preferred habitat (savanna, generalist, or forest). There was an increase in all structural parameters assessed in all vegetation types, with recruitment and gains in basal area and biomass greater than mortality and losses. Thus, there were net gains between the first and final inventories in density (OC: 3.4–22.9%; TC: 1.8–12.6%; DW: 0.2–8.3%), in basal area (OC: 8.3–18.2%; TC: 2–12.7%; DW: 2.3–8.9%), and in biomass (OC: 10.6–16.4%; TC: 1–12%; DW: 5.2–18.7%). Furthermore, all vegetation types also experienced net gains in forest and generalist species relative to savanna species. A decline in recruitment of savanna species was a likely consequence of vegetation encroachment and environmental changes. Our results indicate, for the first time based on quantitative and standardized multi-site temporal data, that concerted structural changes caused by vegetation encroachment are occurring at the ecotone between the two largest biomes in Brazil.

Species distribution models (SDMs) are widely used in ecology and conservation. Presence-only SDMs such as MaxEnt frequently use natural history collections (NHCs) as occurrence data, given their huge numbers and accessibility. NHCs are often spatially biased which may generate inaccuracies in SDMs. Here, we test how the distribution of NHCs and MaxEnt predictions relates to a spatial abundance model, based on a large plot dataset for Amazonian tree species, using inverse distance weighting (IDW). We also propose a new pipeline to deal with inconsistencies in NHCs and to limit the area of occupancy of the species. We found a significant but weak positive relationship between the distribution of NHCs and IDW for 66% of the species. The relationship between SDMs and IDW was also significant but weakly positive for 95% of the species, and sensitivity for both analyses was high. Furthermore, the pipeline removed half of the NHCs records. Presence-only SDM applications should consider this limitation, especially for large biodiversity assessments projects, when they are automatically generated without subsequent checking. Our pipeline provides a conservative estimate of a species' area of occupancy, within an area slightly larger than its extent of occurrence, compatible to e.g. IUCN red list assessments.

Tropical forests play a key role in regulating the terrestrial carbon cycle and climate change by storing a large amount of carbon. Yet, there is considerable uncertainty about the amount and spatial variation of aboveground biomass (AGB), especially in the relatively less studied African tropical forests. In this study, we explore the local-scale variation and determinants of plot-level AGB, between and within two types of forests, the Celtis and Manilkara forests, growing under the same climate but on different geological substrates in the northern Republic of Congo. In each forest site, all trees ≥10 cm diameter were censured in 36 × 1-ha plots and we measured tree height and crown size using a subsample of 18 × 1-ha of these plots. We developed height-diameter and crown-diameter allometric relationships and tested whether they differed between the two sites. For each 1-ha plot, we further estimated the AGB and calculated structural attributes (stem density and basal area), composition attributes (wood density) and architectural attributes (tree height and crown size), the latter being derived from site-specific allometric relationships. We found strong between-site differences in height-diameter and crown-diameter allometries. For a given diameter, trees were taller in the Celtis forest while they had larger crown in the Manilkara forest. Similar trends were found for the sixteen species present in both forest sites, suggesting an environmental control of tree allometry. Although there were some between-site differences in forest structure, composition and architecture, we did not detect any significant difference in mean AGB between the Celtis and the Manilkara forests. The AGB variation was related to the heterogeneous distribution of large trees, and influenced by basal area, height and crown dimensions, and to a lesser extent wood density. These forest attributes have strong practical implications on emerging remote-sensing technologies for carbon monitoring in tropical forests.

Amazon Basin forest soils contain considerable soil organic carbon stocks; however, the contribution of soil pyrogenic carbon (PyC) to the total is unknown. PyC is derived from local fires (historical and modern) and external inputs via aeolian deposition. To establish an initial estimate of PyC stocks in non-terra preta forest with no known history of fire, to assess site and vertical variability, as well as to determine optimal sampling design, we sampled 37 one hectare forest plots in the Amazon Basin and analysed PyC via hydrogen pyrolysis of three individual samples per plot and of bulked samples to 200 cm depth. Using our data and published total organic carbon stocks, we present the first field-based estimate of total PyC stock for the Amazon Basin of 1.10 Pg over 0–30 cm soil depth, and 2.76 Pg over 0–100 cm soil depth. This is up to 20 times higher than previously assumed. Three individual samples per 1 ha are sufficient to capture the site variability of PyC in our plots. PyC showed significant, large-scale variability among plots. To capture 50% of the PyC in 200 cm soil profiles, soil must be sampled to a depth of at least 71 cm. PyC represents a significant (11%) portion of total organic carbon in soil profiles 0–200 cm depth. This finding highlights the potentially important role that historical fire has played in modifying soil C stocks. Our data suggest that PyC is an important carbon pool for long-term storage, involved in millennial scale biogeochemical cycling, particularly in the subsurface soil.

The Amazon region holds a wide variety of ethnic groups and microclimates, enabling different interactions between humans and environment. To better understand the evolution of this region, ancient remains need to be analysed by all possible means. In this context, the study of natural and/or anthropogenic fires through the analysis of carbonized remains can give information on past climate, species diversity, and human intervention in forests and landscapes. In the present work, we undertook an anthracological analysis along with the 14C dating of charcoal fragments using accelerator mass spectrometry (AMS). Charcoal samples from forest soils collected from seven different locations in the Amazon Basin were taxonomically classified and dated. Out of the 16 groups of charcoal fragments identified, five contained more than one taxonomic type, with the Fabaceae, Combretaceae and Sapotaceae families having the highest frequencies. 14C charcoal dates span ∼6000 years (from 6876 to 365 yr BP) among different families, with the most significant variation observed for two fragments from the same sampling location (spanning 4000 14C yr). Some sample sets resulted in up to five different families. These findings demonstrate the importance of the association between anthracological identification and radiocarbon dating in the reconstruction of paleo-forest composition and fire history.

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

Amazon forests are fire-sensitive ecosystems and consequently fires affect forest structure and composition. For instance, the legacy of past fire regimes may persist through some species and traits that are found due to past fires. In this study, we tested for relationships between functional traits that are classically presented as the main components of plant ecological strategies and environmental filters related to climate and historical fires among permanent mature forest plots across the range of local and regional environmental gradients that occur in Amazonia. We used percentage surface soil pyrogenic carbon (PyC), a recalcitrant form of carbon that can persist for millennia in soils, as a novel indicator of historical fire in old-growth forests. Five out of the nine functional traits evaluated across all 378 species were correlated with some environmental variables. Although there is more PyC in Amazonian soils than previously reported, the percentage soil PyC indicated no detectable legacy effect of past fires on contemporary functional composition. More species with dry diaspores were found in drier and hotter environments. We also found higher wood density in trees from higher temperature sites. If Amazon forest past burnings were local and without distinguishable attributes of a widespread fire regime, then impacts on biodiversity would have been small and heterogeneous. Alternatively, sufficient time may have passed since the last fire to allow for species replacement. Regardless, as we failed to detect any impact of past fire on present forest functional composition, if our plots are representative then it suggests that mature Amazon forests lack a compositional legacy of past fire.

Synthetic Aperture Radar (SAR) signals respond to the interactions of microwaves with vegetation canopy scatterers that collectively characterise forest structure. The sensitivity of S-band (7.5-15 cm) backscatter to the different forest types (broadleaved, needleleaved) with varying aboveground biomass (AGB) across temperate (mixed, needleleaved) and tropical (broadleaved, woody savanna, secondary) forests is less well understood. In this study, Michigan Microwave Canopy Scattering (MIMICS-I) radiative transfer model simulations showed strong volume scattering returns from S-band SAR for broadleaved canopies caused by ground/trunk interactions. A general relationship between AirSAR S-band measurements and MIMICS-I simulated radar backscatter with forest AGB up to nearly 100 t/ha in broadleaved forest in the UK was found. Simulated S-band backscatter-biomass relationships suggest increasing backscatter sensitivity to forest biomass with a saturation level close to 100 t/ha and errors between 37 t/ha and 44 t/ha for HV and VV polarisations for tropical ecosystems. In the near future, satellite SAR-derived forest biomass from P-band BIOMASS mission and L-band ALOS-2 PALSAR-2 in combination with S-band UK NovaSAR-S and the joint NASA-ISRO NISAR sensors will provide better quantification of large-scale forest AGB at varying sensitivity levels across primary and secondary forests and woody savannas.

Within the tropics, the species richness of tree communities is strongly and positively associated with precipitation. Previous research has suggested that this macroecological pattern is driven by the negative effect of water-stress on the physiological processes of most tree species. This implies that the range limits of taxa are defined by their ability to occur under dry conditions, and thus in terms of species distributions predicts a nested pattern of taxa distribution from wet to dry areas. However, this ‘dry-tolerance’ hypothesis has yet to be adequately tested at large spatial and taxonomic scales. Here, using a dataset of 531 inventory plots of closed canopy forest distributed across the western Neotropics we investigated how precipitation, evaluated both as mean annual precipitation and as the maximum climatological water deficit, influences the distribution of tropical tree species, genera and families. We find that the distributions of tree taxa are indeed nested along precipitation gradients in the western Neotropics. Taxa tolerant to seasonal drought are disproportionally widespread across the precipitation gradient, with most reaching even the wettest climates sampled; however, most taxa analysed are restricted to wet areas. Our results suggest that the ‘dry tolerance' hypothesis has broad applicability in the world's most species-rich forests. In addition, the large number of species restricted to wetter conditions strongly indicates that an increased frequency of drought could severely threaten biodiversity in this region. Overall, this study establishes a baseline for exploring how tropical forest tree composition may change in response to current and future environmental changes in this region.

©2016. American Geophysical Union. All Rights Reserved. The Amazon Basin has experienced more variable climate over the last decade, with a severe and widespread drought in 2005 causing large basin-wide losses of biomass. A drought of similar climatological magnitude occurred again in 2010; however, there has been no basin-wide ground-based evaluation of effects on vegetation. We examine to what extent the 2010 drought affected forest dynamics using ground-based observations of mortality and growth from an extensive forest plot network. We find that during the 2010 drought interval, forests did not gain biomass (net change: −0.43 Mg ha−1, confidence interval (CI): −1.11, 0.19, n = 97), regardless of whether forests experienced precipitation deficit anomalies. This contrasted with a long-term biomass sink during the baseline pre-2010 drought period (1998 to pre-2010) of 1.33 Mg ha−1yr−1(CI: 0.90, 1.74, p < 0.01). The resulting net impact of the 2010 drought (i.e. reversal of the baseline net sink) was −1.95 Mg ha−1yr−1(CI:−2.77, −1.18; p < 0.001). This net biomass impact was driven by an increase in biomass mortality (1.45 Mg ha−1yr−1CI: 0.66, 2.25, p < 0.001) and a decline in biomass productivity (−0.50 Mg ha−1yr−1, CI:−0.78, −0.31; p < 0.001). Surprisingly, the magnitude of the losses through tree mortality was unrelated to estimated local precipitation anomalies and was independent of estimated local pre-2010 drought history. Thus, there was no evidence that pre-2010 droughts compounded the effects of the 2010 drought. We detected a systematic basin-wide impact of the 2010 drought on tree growth rates across Amazonia, which was related to the strength of the moisture deficit. This impact differed from the drought event in 2005 which did not affect productivity. Based on these ground data, live biomass in trees and corresponding estimates of live biomass in lianas and roots, we estimate that intact forests in Amazonia were carbon neutral in 2010 (−0.07 Pg C yr−1CI:−0.42, 0.23), consistent with results from an independent analysis of airborne estimates of land-atmospheric fluxes during 2010. Relative to the long-term mean, the 2010 drought resulted in a reduction in biomass carbon uptake of 1.1 Pg C, compared to 1.6 Pg C for the 2005 event.

Accurate archaeological and palaeoenvironmental reconstructions using phytoliths relies on the study of modern reference material. In eastern Acre, Brazil, we examined whether the five most common forest types present today were able to be differentiated by their soil phytolith assemblages, and thus provide analogues with which to compare palaeoecological assemblages from pre-Columbian earthwork sites in the region. Surface soils and vegetation from dense humid evergreen forest, dense humid evergreen forest with high palm abundance, palm forest, bamboo forest and fluvial forest were sampled and their phytoliths analysed. Relative phytolith frequencies were statistically compared using Principal Components Analyses (PCAs). We found the major differences in species composition to be well-represented by the phytolith assemblages as all forest types, apart from the two sub-types of dense humid evergreen forest, could be differentiated. Larger phytoliths from the sand fraction were found to be more ecologically diagnostic than those from the silt fraction. The surface soil phytolith assemblages we analysed can therefore be used as analogues to improve the accuracy of archaeological and palaeoecological reconstructions in the region.

Amazon forests, which store ∼ 50% of tropical forest carbon and play a vital role in global water, energy, and carbon cycling, are predicted to experience both longer and more intense dry seasons by the end of the 21st century. However, the climate sensitivity of this ecosystem remains uncertain: several studies have predicted large-scale die-back of the Amazon, whereas several more recent studies predict that the biome will remain largely intact. Combining remote-sensing and ground-based observations with a size- and age-structured terrestrial ecosystem model, we explore the sensitivity and ecological resilience of these forests to changes in climate. We demonstrate that water stress operating at the scale of individual plants, combined with spatial variation in soil texture, explains observed patterns of variation in ecosystem biomass, composition, and dynamics across the region, and strongly influences the ecosystem's resilience to changes in dry season length. Specifically, our analysis suggests that in contrast to existing predictions of either stability or catastrophic biomass loss, the Amazon forest's response to a drying regional climate is likely to be an immediate, graded, heterogeneous transition from high-biomass moist forests to transitional dry forests and woody savannah-like states. Fire, logging, and other anthropogenic disturbances may, however, exacerbate these climate change-induced ecosystem transitions.

Empirical evidence and modelling both suggest that global changes may lead to an increased dominance of lianas and thus to an increased prevalence of liana-infested forest formations in tropical forests. The implications for tropical forest structure and the carbon cycle remain poorly understood. We studied the ecological processes underpinning the structure and dynamics of a liana-infested forest in French Guiana, using a combination of long-term surveys (tree, liana, seedling and litterfall), soil chemical analyses and remote-sensing approaches (LiDAR and Landsat). At stand scale and for adult trees, the liana-infested forest had higher growth, recruitment and mortality rates than the neighbouring high-canopy forest. Both total seedling density and tree seedling recruitment were lower in the liana-infested forest. Stand scale above-ground biomass of the liana-infested forest was 58% lower than in the high-canopy forest. Above-ground net primary productivity (ANPP) was comparable in the liana-infested and high-canopy forests. However, due to more abundant leaf production, the relative contribution of fast turnover carbon pools to ANPP was larger in the liana-infested forest and the carbon residence time was half that of the high-canopy forest. Although soils of the liana-infested forest were richer in nutrients, soil elemental ratios suggest that liana-infested forest and high-canopy forest soils both derive from the same geological substrate. The higher nutrient concentration in the liana-infested forest may therefore be the result of a release of nutrients from vegetation after a forest blowdown. Using small-footprint LiDAR campaigns, we show that the overall extent of the liana-infested forest has remained stable from 2007 to 2012 but about 10% of the forest area changed in forest cover type. Landsat optical imagery confirms the liana-infested forest presence in the landscape for at least 25 years. Synthesis. Because persistently high rates of liana infestation are maintained by the fast dynamics of the liana-infested forest, liana-infested forests here appear to be the result of an arrested tropical forest succession. If the prevalence of such arrested succession forests were to increase in the future, this would have important implications for the carbon sink potential of Amazonian forests. Because persistently high rates of liana infestation are maintained by the fast dynamics of the liana-infested forest, liana-infested forests here appear to be the result of an arrested tropical forest succession. If the prevalence of such arrested succession forests were to increase in the future, this would have important implications for the carbon sink potential of Amazonian forests. Journal of Ecology

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.

The loss of biodiversity in transitional forests between the Cerrado and Amazonia, the two largest neotropical phytogeographic domains, is an issue of great concern. This extensive region is located within the 'arc of deforestation' zone where tropical forests are being lost at the fastest rate on the planet, but floristic diversity and variation among forests here is still poorly understood. We aimed to characterize the floristic composition of forests in this zone and explored the degree and drivers of differentiation within and across Araguaia and Xingu watersheds. In 10 sites we identified all trees with diameter ≥10 cm; these totaled 4944 individuals in 257 species, 107 genera and 52 families. We evaluated the data for multivariate variation using TWINSPAN and DCA to understand the species distribution among sites. There was a larger contribution from the Amazonian flora (169 species) than that of the Cerrado (109) to the transitional forests. Remarkably, 142 species (55%) were restricted to only one sampling site, while 29 species (>16%) are endemic to Brazil, suggesting potentially large loss of species and unique forest communities with the loss and fragmentation of large areas. Our results also suggest that watersheds may be a critical factor driving species distribution among forests in the Amazonian-Cerrado transition zone, and quantifying their role can provide powerful insight into devising better conservation strategies for the remaining forests.

Developing and improving methods to monitor forest carbon in space and time is a timely challenge, especially for tropical forests. The next European Space Agency Earth Explorer Core Mission BIOMASS will collect synthetic aperture radar (SAR) data globally from employing a multiple baseline orbit during the initial phase of its lifetime. These data will be used for tomographic SAR (TomoSAR) processing, with a vertical resolution of about 20 m, a resolution sufficient to decompose the backscatter signal into two to three layers for most closed-canopy tropical forests. A recent study, conducted in the Paracou site, French Guiana, has already shown that TomoSAR significantly improves the retrieval of forest aboveground biomass (AGB) in a high biomass forest, with an error of only 10% at 1.5-ha resolution. However, the degree to which this TomoSAR approach can be transferred from one site to another has not been assessed. We test this approach at the Nouragues site in central French Guiana (ca 100 km away from Paracou), and develop a method to retrieve the top-of-canopy height from TomoSAR. We found a high correlation between the backscatter signal and AGB in the upper canopy layer (i.e. 20-40 m), while lower layers only showed poor correlations. The relationship between AGB and TomoSAR data was found to be highly similar for forests at Nouragues and Paracou. Cross validation using training plots from Nouragues and validation plots from Paracou, and vice versa, gave an error of 16-18% of AGB using 1-ha plots. Finally, using a high-resolution LiDAR canopy model as a reference, we showed that TomoSAR has the potential to retrieve the top-of-canopy height with an error to within 2.5 m. Our analyses show that the TomoSAR-AGB retrieval method is accurate even in hilly and high-biomass forest areas and suggest that our approach may be generalizable to other study sites, having a canopy taller than 30 m. These results have strong implications for the tomographic phase of the BIOMASS spaceborne mission.

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 (DGVMs). 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 DGVMs. 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, DGVMs 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 DGVMs.

Photosynthesis/nutrient relationships of proximally growing forest and savanna trees were determined in an ecotonal region of Cameroon (Africa). Although area-based foliar N concentrations were typically lower for savanna trees, there was no difference in photosynthetic rates between the two vegetation formation types. Opposite to N, area-based P concentrations were—on average—slightly lower for forest trees; a dependency of photosynthetic characteristics on foliar P was only evident for savanna trees. Thus savanna trees use N more efficiently than their forest counterparts, but only in the presence of relatively high foliar P. Along with some other recent studies, these results suggest that both N and P are important modulators of woody tropical plant photosynthetic capacities, influencing photosynthetic metabolism in different ways that are also biome specific. Attempts to find simple unifying equations to describe woody tropical vegetation photosynthesis-nutrient relationships are likely to meet with failure, with ecophysiological distinctions between forest and savanna requiring acknowledgement.

We compared the diversity and species composition and the structure of the vegetation of three distinct Cerrado phytophysiognomies (Cerradão, Dense Cerrado, and Typical Cerrado) in the Cerrado–Amazon transition, Mato Grosso (Brazil). Species richness (observed and estimated) in the Cerradão and Dense Cerrado was higher than that recorded in the Typical Cerrado. Species diversity, based on a Rényi profile, was highest in the Dense Cerrado, in comparison with the other phytophysiognomies. We recorded a higher number of exclusive species in the Cerradão and a greater similarity (Morisita and Sørensen indices) between this vegetation type and the Dense Cerrado. While individuals were tallest in the Cerradão and Dense Cerrado and lowest in the Typical Cerrado, there was no difference among phytophysiognomies in mean diameter. A gradient in decreasing species richness and diversity (hypothesis 1) and vegetation vertical structure (hypothesis 3) was expected for the Cerradão–Dense Cerrado–Typical Cerrado; however, neither hypothesis was supported by the results. The Cerradão and Dense Cerrado were most similar in species composition not confirming hypothesis 2, which predicted that the two savanna vegetation types (Dense Cerrado and Typical Cerrado) were more similar to one another than either is to the woodland (Cerradão). Overall, the similarities among the three study communities depended on the type of parameter analyzed. While the species richness and the vertical and structure of the vegetation of the Cerradão and Dense Cerrado are closely similar, the Cerradão and Typical Cerrado are more similar in their species diversity. With regard to the floristic composition, Dense Cerrado occupies an intermediate position between Cerradão and Typical Cerrado.

In 2005 and 2010 the Amazon basin experienced two strong droughts, driven by shifts in the tropical hydrological regime possibly associated with global climate change, as predicted by some global models. Tree mortality increased after the 2005 drought, and regional atmospheric inversion modelling showed basin-wide decreases in CO2 uptake in 2010 compared with 2011 (ref. 5). But the response of tropical forest carbon cycling to these droughts is not fully understood and there has been no detailed multi-site investigation in situ. Here we use several years of data from a network of thirteen 1-ha forest plots spread throughout South America, where each component of net primary production (NPP), autotrophic respiration and heterotrophic respiration is measured separately, to develop a better mechanistic understanding of the impact of the 2010 drought on the Amazon forest. We find that total NPP remained constant throughout the drought. However, towards the end of the drought, autotrophic respiration, especially in roots and stems, declined significantly compared with measurements in 2009 made in the absence of drought, with extended decreases in autotrophic respiration in the three driest plots. In the year after the drought, total NPP remained constant but the allocation of carbon shifted towards canopy NPP and away from fine-root NPP. Both leaf-level and plot-level measurements indicate that severe drought suppresses photosynthesis. Scaling these measurements to the entire Amazon basin with rainfall data, we estimate that drought suppressed Amazon-wide photosynthesis in 2010 by 0.38 petagrams of carbon (0.23-0.53 petagrams of carbon). Overall, we find that during this drought, instead of reducing total NPP, trees prioritized growth by reducing autotrophic respiration that was unrelated to growth. This suggests that trees decrease investment in tissue maintenance and defence, in line with eco-evolutionary theories that trees are competitively disadvantaged in the absence of growth. We propose that weakened maintenance and defence investment may, in turn, cause the increase in post-drought tree mortality observed at our plots.

Premise of research. This represents one of the first studies of the ecology, diversity, and structure of campos de murundus termite savannas in the vast seasonal wetlands of southern Amazonia. We aimed to improve understanding of this threatened system by assessing species richness, abundance, and co-occurrence among trees and herbs of murundus (earth mounds), investigating the environmental and biological mechanisms underlying these patterns, and discussing implications for biodiversity conservation. Methodology. We identified every tree, shrub, subshrub, and herb on 373 murundus across 11 ha at Araguaia State Park, southern Amazonia. We constructed species abundance distributions of trees and herbs, assessed best-fit models, and tested for nonrandom patterns of species co-occurrence using checkerboard scores. Using detrended correspondence analysis (DCA), we assessed the affinities among tree species and their positions in murundus. Pivotal results. A total of 166 species, 123 genera, and 49 families occupied the murundus. The species abundance distribution of trees followed a lognormal distribution, whereas that of herbs was best described by a Mandelbrot distribution. Observed C-score indices for trees and herbs were significantly larger than expected by chance, indicating nonrandom distributions and species segregation among murundus. DCA revealed a strong gradient in species occurrence within murundus, suggesting that internal structuring may be hydrologically based (e.g. variation in mound microrelief). Conclusions. Environmental (e.g. flooding) and biological (e.g. competition between plants) factors are important for controlling the occurrence of tree and herb species on the murundus. The murundus function as critical bases for the maintenance of species diversity in this extensive floodplain, thereby deserving recognition among ecosystems with high conservation priorities.

Sampling along a precipitation gradient in tropical South America extending from ca. 0.8 to 2.0 m ag-1, savanna soils had consistently lower exchangeable cation concentrations and higher C/N ratios than nearby forest plots. These soil differences were also reflected in canopy averaged leaf traits with savanna trees typically having higher leaf mass per unit area but lower mass-based nitrogen (Nm) and potassium (Km). Both Nm and Km also increased with declining mean annual precipitation (PA), but most area-based leaf traits such as leaf photosynthetic capacity showed no systematic variation with PA or vegetation type. Despite this invariance, when taken in conjunction with other measures such as mean canopy height, area-based soil exchangeable potassium content, [K]sa , proved to be an excellent predictor of several photosynthetic properties (including 13C isotope discrimination). Moreover, when considered in a multivariate context with PA and soil plant available water storage capacity (θP) as covariates, [K]sa also proved to be an excellent predictor of stand-level canopy area, providing drastically improved fits as compared to models considering just PA and/or θP. Neither calcium, nor magnesium, nor soil pH could substitute for potassium when tested as alternative model predictors (ΔAIC > 10). Nor for any model could simple soil texture metrics such as sand or clay content substitute for either [K]sa or θP. Taken in conjunction with recent work in Africa and the forests of the Amazon Basin, this suggests-in combination with some newly conceptualised interacting effects of PA and θP also presented here-a critical role for potassium as a modulator of tropical vegetation structure and function.

Estimates of extinction risk for Amazonian plant and animal species are rare and not often incorporated into land-use policy and conservation planning. We overlay spatial distribution models with historical and projected deforestation to show that at least 36% and up to 57% of all Amazonian tree species are likely to qualify as globally threatened under International Union for Conservation of Nature (IUCN) Red List criteria. If confirmed, these results would increase the number of threatened plant species on Earth by 22%. We show that the trends observed in Amazonia apply to trees throughout the tropics, and we predict that most of the world's >40,000 tropical tree species now qualify as globally threatened. A gap analysis suggests that existing Amazonian protected areas and indigenous territories will protect viable populations of most threatened species if these areas suffer no further degradation, highlighting the key roles that protected areas, indigenous peoples, and improved governance can play in preventing large-scale extinctions in the tropics in this century.

Variations in leaf mass per unit area (Ma) and foliar concentrations of N, P, C, K, Mg and Ca were determined for 365 trees growing in 23 plots along a West African precipitation gradient ranging from 0.29 to 1.62m a-1. Contrary to previous studies, no marked increase in Ma with declining precipitation was observed, but savanna tree foliar [N] tended to be higher at the drier sites (mass basis). Generally, Ma was slightly higher and [N] slightly lower for forest vs savanna trees with most of this difference attributable to differences in soil chemistry. No systematic variations in [P], [Mg] and [Ca] with precipitation or between trees of forest vs savanna stands were observed. We did, however, find a marked increase in foliar [K] of savanna trees as precipitation declined, with savanna trees also having a significantly lower [K] than those of nearby forest. These differences were not related to differences in soil nutrient status and were accompanied by systematic changes in [C] of opposite sign. We suggest an important but as yet unidentified role for K in the adaption of savanna species to periods of limited water availability; with foliar [K] being also an important factor differentiating tree species adapted to forest vs savanna soils within the 'zone of transition' of Western Africa.

While Amazonian forests are extraordinarily diverse, the abundance of trees is skewed strongly towards relatively few 'hyperdominant' species. In addition to their diversity, Amazonian trees are a key component of the global carbon cycle, assimilating and storing more carbon than any other ecosystem on Earth. Here we ask, using a unique data set of 530 forest plots, if the functions of storing and producing woody carbon are concentrated in a small number of tree species, whether the most abundant species also dominate carbon cycling, and whether dominant species are characterized by specific functional traits. We find that dominance of forest function is even more concentrated in a few species than is dominance of tree abundance, with only ≈1% of Amazon tree species responsible for 50% of carbon storage and productivity. Although those species that contribute most to biomass and productivity are often abundant, species maximum size is also influential, while the identity and ranking of dominant species varies by function and by region.

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.

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 (PDss), the mean pairwise phylogenetic distance between species (MPD), the mean nearest taxon distance (MNTD) and their equivalents standardized for species richness (ses.PDss, 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: PDss 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 PDss and the lowest MNTD. We find that the youngest and oldest substrates (the latter on the Brazilian and Guiana Shields) have the highest ses.PDss 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.PDss and ses.MNTD reflect greater lineage diversity in communities. We suggest that high ses.PDss 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 PDss, ses.PDss and ses.MNTD may be the most useful diversity metrics for setting large-scale conservation priorities.

Fire can change the species composition, diversity, and structure of savanna vegetation, thus altering growth and mortality rates. Such changes in the woody vegetation of burned savanna forest were evaluated over four years in comparison to unburned savanna forest. All woody plants with a diameter at breast height > 10 cm were measured in 100 permanent plots. Six months later, 38 of these plots were burned. Three and a half years later, all surviving individuals were re-sampled. Species richness, diversity, and the number of individuals did not change in the burned plots, although they had significantly higher (p < 0.05) increases in basal area and mortality rates (5.1% year1) than the unburned plots (3.0% year-1). Tachigali vulgaris had the greatest post-fire increase in basal area (53%). The results indicate that fire alters the dynamics and structure of the savanna forest, excluding the less fire-tolerant species and smaller individuals (< 15cm). Tachigali vulgaris is a key species for the recovery of savanna forest biomass due to its considerable post-fire gains in basal area, at least over the short term due to its short life cycle. It follows that frequent burning of savanna forest would result in a marked change in the species composition and structure of its woody vegetation.

Recent analyses of Amazon runoff and gridded precipitation data suggest an intensification of the hydrological cycle over the past few decades in the following sense: wet season precipitation and peak river runoff (since ∼1980) as well as annual mean precipitation (since ∼1990) have increased, while dry season precipitation and minimum runoff have slightly decreased. There has also been an increase in the frequency of anomalously severe floods and droughts. To provide context for the special issue on Amazonia and its forests in a warming climate we expand here on these analyses. The contrasting recent changes in wet and dry season precipitation have continued and are generally consistent with changes in catchment-level peak and minimum river runoff as well as a positive trend of water vapor inflow into the basin. Consistent with the river records, the increased vapor inflow is concentrated to the wet season. Temperature has been rising by 0.7°C since 1980 with more pronounced warming during dry months. Suggestions for the cause of the observed changes of the hydrological cycle come from patterns in tropical sea surface temperatures (SSTs). Tropical and North Atlantic SSTs have increased rapidly and steadily since 1990, while Pacific SSTs have shifted during the 1990s from a positive Pacific Decadal Oscillation (PDO) phase with warm eastern Pacific temperatures to a negative phase with cold eastern Pacific temperatures. These SST conditions have been shown to be associated with an increase in precipitation over most of the Amazon except the south and southwest. If ongoing changes continue, we expect forests to continue to thrive in those regions where there is an increase in precipitation with the exception of floodplain forests. An increase in flood pulse height and duration could lead to increased mortality at higher levels of the floodplain and, over the long term, to a lateral shift of the zonally stratified floodplain forest communities. Negative effects on forests are mainly expected in the southwest and south, which have become slightly drier and hotter, consistent with tree mortality trends observed at the RAINFOR Amazon forest plot network established in the early 1980s consisting of approximately 150 regularly censused 1ha plots in intact forests located across the whole basin.

Through interpretations of remote-sensing data and/or theoretical propositions, the idea that forest and savanna represent "alternative stable states" is gaining increasing acceptance. Filling an observational gap, we present detailed stratified floristic and structural analyses for forest and savanna stands located mostly within zones of transition (where both vegetation types occur in close proximity) in Africa, South America and Australia. Woody plant leaf area index variation was related to tree canopy cover in a similar way for both savanna and forest with substantial overlap between the two vegetation types. As total woody plant canopy cover increased, so did the relative contribution of middle and lower strata of woody vegetation. Herbaceous layer cover declined as woody cover increased. This pattern of understorey grasses and herbs progressively replaced by shrubs as the canopy closes over was found for both savanna and forests and on all continents. Thus, once subordinate woody canopy layers are taken into account, a less marked transition in woody plant cover across the savanna-forest-species discontinuum is observed compared to that inferred when trees of a basal diameter > 0.1 m are considered in isolation. This is especially the case for shrub-dominated savannas and in taller savannas approaching canopy closure. An increased contribution of forest species to the total subordinate cover is also observed as savanna stand canopy closure occurs. Despite similarities in canopy-cover characteristics, woody vegetation in Africa and Australia attained greater heights and stored a greater amount of above-ground biomass than in South America. Up to three times as much above-ground biomass is stored in forests compared to savannas under equivalent climatic conditions. Savanna-forest transition zones were also found to typically occur at higher precipitation regimes for South America than for Africa. Nevertheless, consistent across all three continents coexistence was found to be confined to a well-defined edaphic-climate envelope with soil and climate the key determinants of the relative location of forest and savanna stands. Moreover, when considered in conjunction with the appropriate water availability metrics, it emerges that soil exchangeable cations exert considerable control on woody canopy-cover extent as measured in our pan-continental (forest + savanna) data set. Taken together these observations do not lend support to the notion of alternate stable states mediated through fire feedbacks as the prime force shaping the distribution of the two dominant vegetation types of the tropical lands.

Variations in the carbon isotopic composition of soil organic matter (SOM) in bulk and fractionated samples were used to assess the influence of C3 and C4 vegetation on SOM dynamics in semi-natural tropical ecosystems sampled along a precipitation gradient in West Africa. Differential patterns in SOM dynamics in C3/C4 mixed ecosystems occurred at various spatial scales. Relative changes in C / N ratios between two contrasting SOM fractions were used to evaluate potential site-scale differences in SOM dynamics between C3- and C4-dominated locations. These differences were strongly controlled by soil texture across the precipitation gradient, with a function driven by bulk δ13C and sand content explaining 0.63 of the observed variability. The variation of δ13C with soil depth indicated a greater accumulation of C3-derived carbon with increasing precipitation, with this trend also being strongly dependant on soil characteristics. The influence of vegetation thickening on SOM dynamics was also assessed in two adjacent, but structurally contrasting, transitional ecosystems occurring on comparable soils to minimise the confounding effects posed by climatic and edaphic factors. Radiocarbon analyses of sand-size aggregates yielded relatively short mean residence times (τ) even in deep soil layers, while the most stable SOM fraction associated with silt and clay exhibited shorter τ in the savanna woodland than in the neighbouring forest stand. These results, together with the vertical variation observed in δ13C values, strongly suggest that both ecosystems are undergoing a rapid transition towards denser closed canopy formations. However, vegetation thickening varied in intensity at each site and exerted contrasting effects on SOM dynamics. This study shows that the interdependence between biotic and abiotic factors ultimately determine whether SOM dynamics of C3- and C4-derived vegetation are at variance in ecosystems where both vegetation types coexist. The results highlight the far-reaching implications that vegetation thickening may have for the stability of deep SOM. Â

In recent years, LiDAR technology has provided accurate forest aboveground biomass (AGB) maps in several forest ecosystems, including tropical forests. However, its ability to accurately map forest AGB changes in high-biomass tropical forests has seldom been investigated. Here, we assess the ability of repeated LiDAR acquisitions to map AGB stocks and changes in an old-growth Neotropical forest of French Guiana. Using two similar aerial small-footprint LiDAR campaigns over a four year interval, spanning ca. 20km2, and concomitant ground sampling, we constructed a model relating median canopy height and AGB at a 0.25-ha and 1-ha resolution. This model had an error of 14% at a 1-ha resolution (RSE=54.7Mgha-1) and of 23% at a 0.25-ha resolution (RSE=86.5Mgha-1). This uncertainty is comparable with values previously reported in other tropical forests and confirms that aerial LiDAR is an efficient technology for AGB mapping in high-biomass tropical forests. Our map predicts a mean AGB of 340Mgha-1 within the landscape. We also created an AGB change map, and compared it with ground-based AGB change estimates. The correlation was weak but significant only at the 0.25-ha resolution. One interpretation is that large natural tree-fall gaps that drive AGB changes in a naturally regenerating forest can be picked up at fine spatial scale but are veiled at coarser spatial resolution. Overall, both field-based and LiDAR-based estimates did not reveal a detectable increase in AGB stock over the study period, a trend observed in almost all forest types of our study area. Small footprint LiDAR is a powerful tool to dissect the fine-scale variability of AGB and to detect the main ecological controls underpinning forest biomass variability both in space and time.

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.

Forest inventory plots are widely used to estimate biomass carbon storage and its change over time. While there has been much debate and exploration of the analytical methods for calculating biomass, the methods used to determine rates of wood production have not been evaluated to the same degree. This affects assessment of ecosystem fluxes and may have wider implications if inventory data are used to parameterise biospheric models, or scaled to large areas in assessments of carbon sequestration. Here we use a dataset of 35 long-term Amazonian forest inventory plots to test different methods of calculating wood production rates. These address potential biases associated with three issues that routinely impact the interpretation of tree measurement data: (1) changes in the point of measurement (POM) of stem diameter as trees grow over time; (2) unequal length of time between censuses; and (3) the treatment of trees that pass the minimum diameter threshold ("recruits"). We derive corrections that control for changing POM height, that account for the unobserved growth of trees that die within census intervals, and that explore different assumptions regarding the growth of recruits during the previous census interval. For our dataset we find that annual aboveground coarse wood production (AGWP; in Mgha-1year-1 of dry matter) is underestimated on average by 9.2% if corrections are not made to control for changes in POM height. Failure to control for the length of sampling intervals results in a mean underestimation of 2.7% in annual AGWP in our plots for a mean interval length of 3.6years. Different methods for treating recruits result in mean differences of up to 8.1% in AGWP. In general, the greater the length of time a plot is sampled for and the greater the time elapsed between censuses, the greater the tendency to underestimate wood production. We recommend that POM changes, census interval length, and the contribution of recruits should all be accounted for when estimating productivity rates, and suggest methods for doing this. © 2014 Elsevier B.V.

Forest inventory studies in the Amazon indicate a large terrestrial carbon sink. However, field plots may fail to represent forest mortality processes at landscape-scales of tropical forests. Here we characterize the frequency distribution of disturbance events in natural forests from 0.01 ha to 2,651 ha size throughout Amazonia using a novel combination of forest inventory, airborne lidar and satellite remote sensing data. We find that small-scale mortality events are responsible for aboveground biomass losses of ~1.7 Pg C y(-1) over the entire Amazon region. We also find that intermediate-scale disturbances account for losses of ~0.2 Pg C y(-1), and that the largest-scale disturbances as a result of blow-downs only account for losses of ~0.004 Pg C y(-1). Simulation of growth and mortality indicates that even when all carbon losses from intermediate and large-scale disturbances are considered, these are outweighed by the net biomass accumulation by tree growth, supporting the inference of an Amazon carbon sink.

We report above-ground biomass (AGB), basal area, stemdensity and wood mass density estimates from 260 sample plots (mean size: 1.2 ha) in intact closed-canopy tropical forests across 12 African countries. Mean AGB is 395.7 Mg dry mass ha-1 (95% CI: 14.3), substantially higher than Amazonian values, with the Congo Basin and contiguous forest region attaining AGB values (429 Mg ha-1) similar to those of Bornean forests, and significantly greater than East or West African forests. AGB therefore appears generally higher in palaeo- comparedwithneotropical forests.However, mean stem density is low(426±11 stems ha-1 greater than or equal to 100 mm diameter) compared with both Amazonian and Bornean forests (cf. approx. 600) and is the signature structural feature of African tropical forests. While spatial autocorrelation complicates analyses, AGB shows a positive relationship with rainfall in the driest nine months of the year, and an opposite association with the wettest three months of the year; a negative relationship with temperature; positive relationship with clay-rich soils; and negative relationshipswith C:Nratio (suggesting a positive soil phosphorus- AGB relationship), and soil fertility computed as the sum of base cations. The results indicate that AGB is mediated by both climate and soils, and suggest that the AGB of African closed-canopy tropical forests may be particularly sensitive to future precipitation and temperature changes. © 2013 the Authors.

The vast extent of the Amazon Basin has historically restricted the study of its tree communities to the local and regional scales. Here, we provide empirical data on the commonness, rarity, and richness of lowland tree species across the entire Amazon Basin and Guiana Shield (Amazonia), collected in 1170 tree plots in all major forest types. Extrapolations suggest that Amazonia harbors roughly 16,000 tree species, of which just 227 (1.4%) account for half of all trees. Most of these are habitat specialists and only dominant in one or two regions of the basin. We discuss some implications of the finding that a small group of species - less diverse than the North American tree flora - accounts for half of the world's most diverse tree community.

The Amazon basin hosts half the planet's remaining moist tropical forests, but they may be threatened in a warming world. Nevertheless, climate model predictions vary from rapid drying to modest wetting. Here we report that the catchment of the world's largest river is experiencing a substantial wetting trend since approximately 1990. This intensification of the hydrological cycle is concentrated overwhelmingly in the wet season driving progressively greater differences in Amazon peak and minimum flows. The onset of the trend coincides with the onset of an upward trend in tropical Atlantic sea surface temperatures (SST). This positive longer-term correlation contrasts with the short-term, negative response of basin-wide precipitation to positive anomalies in tropical North Atlantic SST, which are driven by temporary shifts in the intertropical convergence zone position. We propose that the Amazon precipitation changes since 1990 are instead related to increasing atmospheric water vapor import from the warming tropical Atlantic. Key PointsIntensification of Amazon Hydrological Cycle since 1990Revealed by both river discharge and precipitation recordsIn parallel onset of tropical Atlantic warming offering explanation ©2013. American Geophysical Union. All Rights Reserved.

Background: There is no generally agreed classification scheme for the many different vegetation formation types occurring in the tropics. This hinders cross-continental comparisons and causes confusion as words such as 'forest' and 'savanna' have different meanings to different people. Tropical vegetation formations are therefore usually imprecisely and/or ambiguously defined in modelling, remote sensing and ecological studies.Aims: to integrate observed variations in tropical vegetation structure and floristic composition into a single classification scheme.Methods: Using structural and floristic measurements made on three continents, discrete tropical vegetation groupings were defined on the basis of overstorey and understorey structure and species compositions by using clustering techniques.Results: Twelve structural groupings were identified based on height and canopy cover of the dominant upper stratum and the extent of lower-strata woody shrub cover and grass cover. Structural classifications did not, however, always agree with those based on floristic composition, especially for plots located in the forest-savanna transition zone. This duality is incorporated into a new tropical vegetation classification scheme.Conclusions: Both floristics and stand structure are important criteria for the meaningful delineation of tropical vegetation formations, especially in the forest/savanna transition zone. A new tropical vegetation classification scheme incorporating this information has been developed. © 2013 Copyright 2013 Botanical Society of Scotland and Taylor & Francis.

The future of tropical forests under global environmental change is uncertain, with biodiversity and carbon stocks at risk if precipitation regimes alter. Here, we assess changes in plant functional composition and biomass in 19 plots from a variety of forest types during two decades of long-term drought in Ghana. We find a consistent increase in dry forest, deciduous, canopy species with intermediate light demand and a concomitant decrease in wet forest, evergreen, sub-canopy and shade-tolerant species. These changes in composition are accompanied by an increase in above-ground biomass. Our results indicate that by altering composition in favour of drought-tolerant species, the biomass stocks of these forests may be more resilient to longer term drought than short-term studies of severe individual droughts suggest.

We summarise the contemporary carbon budget of South America and relate it to its dominant controls: population and economic growth, changes in land use practices and a changing atmospheric environment and climate. Component flux estimate methods we consider sufficiently reliable for this purpose encompass fossil fuel emission inventories, biometric analysis of old-growth rainforests, estimation of carbon release associated with deforestation based on remote sensing and inventories, and agricultural export data. Alternative methods for the estimation of the continental-scale net land to atmosphere CO2 flux, such as atmospheric transport inverse modelling and terrestrial biosphere model predictions, are, we find, hampered by the data paucity, and improved parameterisation and validation exercises are required before reliable estimates can be obtained. From our analysis of available data, we suggest that South America was a net source to the atmosphere during the 1980s (∼ 0.3-0.4 Pg C a&minus;1) and close to neutral (∼ 0.1 Pg C a&minus;1) in the 1990s. During the latter period, carbon uptake in old-growth forests nearly compensated for the carbon release associated with fossil fuel burning and deforestation. Annual mean precipitation over tropical South America as inferred from Amazon River discharge shows a long-term upward trend. Although, over the last decade dry seasons have tended to be drier, with the years 2005 and 2010 in particular experiencing strong droughts. On the other hand, precipitation during the wet seasons also shows an increasing trend. Air temperatures have also increased slightly. Also with increases in atmospheric CO2 concentrations, it is currently unclear what effect these climate changes are having on the forest carbon balance of the region. Current indications are that the forests of the Amazon Basin have acted as a substantial long-term carbon sink, but with the most recent measurements suggesting that this sink may be weakening. Economic development of the tropical regions of the continent is advancing steadily, with exports of agricultural products being an important driver and witnessing a strong upturn over the last decade. © Author(s) 2012.

We examine the influence of climate, soil properties and vegetation characteristics on soil organic carbon (SOC) along a transect of West African ecosystems sampled across a precipitation gradient on contrasting soil types stretching from Ghana (15°N) to Mali (7°N). Our findings derive from a total of 1108 soil cores sampled over 14 permanent plots. The observed pattern in SOC stocks reflects the very different climatic conditions and contrasting soil properties existing along the latitudinal transect. The combined effects of these factors strongly influence vegetation structure. SOC stocks in the first 2 m of soil ranged from 20 Mg C ha -1 for a Sahelian savanna in Mali to over 120 Mg C ha -1 for a transitional forest in Ghana. The degree of interdependence between soil bulk density (SBD) and soil properties is highlighted by the strong negative relationships observed between SBD and SOC (r 2 > 0.84). A simple predictive function capable of encompassing the effect of climate, soil properties and vegetation type on SOC stocks showed that available water and sand content taken together could explain 0.84 and 0.86 of the total variability in SOC stocks observed to 0.3 and 1.0 m depth respectively. Used in combination with a suitable climatic parameter, sand content is a good predictor of SOC stored in highly weathered dry tropical ecosystems with arguably less confounding effects than provided by clay content. There was an increased contribution of resistant SOC to the total SOC pool for lower rainfall soils, this likely being the result of more frequent fire events in the grassier savannas of the more arid regions. This work provides new insights into the mechanisms determining the distribution of carbon storage in tropical soils and should contribute significantly to the development of robust predictive models of biogeochemical cycling and vegetation dynamics in tropical regions. © 2012 Blackwell Publishing Ltd.

Aim: to test the extent to which the vertical structure of tropical forests is determined by environment, forest structure or biogeographical history. Location: Pan-tropical. Methods: Using height and diameter data from 20,497 trees in 112 non-contiguous plots, asymptotic maximum height (H AM) and height-diameter relationships were computed with nonlinear mixed effects (NLME) models to: (1) test for environmental and structural causes of differences among plots, and (2) test if there were continental differences once environment and structure were accounted for; persistence of differences may imply the importance of biogeography for vertical forest structure. NLME analyses for floristic subsets of data (only/excluding Fabaceae and only/excluding Dipterocarpaceae individuals) were used to examine whether family-level patterns revealed biogeographical explanations of cross-continental differences. Results: H AM and allometry were significantly different amongst continents. H AM was greatest in Asian forests (58.3 ± 7.5m, 95% CI), followed by forests in Africa (45.1 ± 2.6m), America (35.8 ± 6.0m) and Australia (35.0 ± 7.4m), and height-diameter relationships varied similarly; for a given diameter, stems were tallest in Asia, followed by Africa, America and Australia. Precipitation seasonality, basal area, stem density, solar radiation and wood density each explained some variation in allometry and H AM yet continental differences persisted even after these were accounted for. Analyses using floristic subsets showed that significant continental differences in H AM and allometry persisted in all cases. Main conclusions: Tree allometry and maximum height are altered by environmental conditions, forest structure and wood density. Yet, even after accounting for these, tropical forest architecture varies significantly from continent to continent. The greater stature of tropical forests in Asia is not directly determined by the dominance of the family Dipterocarpaceae, as on average non-dipterocarps are equally tall. We hypothesise that dominant large-statured families create conditions in which only tall species can compete, thus perpetuating a forest dominated by tall individuals from diverse families. © 2012 Blackwell Publishing Ltd.

We present a critical evaluation of the above-ground biomass (AGB) map of Africa published in this journal by Baccini et al (2008 Environ. Res. Lett. 3 045011). We first test their map against an independent dataset of 1154 scientific inventory plots from 16 African countries, and find only weak correspondence between our field plots and the AGB value given for the surrounding 1km pixel by Baccini et al. Separating our field data using a continental landcover classification suggests that the Baccini et al map underestimates the AGB of forests and woodlands, while overestimating the AGB of savannas and grasslands. Secondly, we compare their map to 216 000 × 0.25ha spaceborne LiDAR footprints. A comparison between Lorey's height (basal-area-weighted average height) derived from the LiDAR data for 1km pixels containing at least five LiDAR footprints again does not support the hypothesis that the Baccini et al map is accurate, and suggests that it significantly underestimates the AGB of higher AGB areas. We conclude that this is due to the unsuitability of some of the field data used by Baccini et al to create their map, and overfitting in their model, resulting in low accuracies outside the small areas from which their field data are drawn. © 2011 IOP Publishing Ltd.

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 &minus;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. © 2011 Author(s).

Satellite L-band synthetic aperture radar backscatter data from 1996 and 2007 (from JERS-1 and ALOS PALSAR respectively), were used with field data collected in 2007 and a back-calibration method to produce biomass maps of a 15000km2 forest-savanna ecotone region of central Cameroon. The relationship between the radar backscatter and aboveground biomass (AGB) was strong (r2=0.86 for ALOS HV to biomass plots, r2=0.95 relating ALOS-derived biomass for 40 suspected unchanged regions to JERS-1 HH). The root mean square error (RMSE) associated with AGB estimation varied from ~25% for AGB100Mgha-1 for the ALOS HV data. Change detection showed a significant loss of AGB over high biomass forests, due to suspected deforestation and degradation, and significant biomass gains along the forest-savanna boundary, particularly in areas of low population density. Analysis of the errors involved showed that radar data can detect changes in broad AGB class in forest-savanna transition areas with an accuracy >95%. However, quantitative assessment of changes in AGB in Mgha-1 at a pixel level will require radar images from sensors with similar characteristics collecting data from the same season over multiple years. © 2011 Elsevier Inc.

Stocks of dead wood or necromass represent an important portion of biomass and nutrients in tropical forests. The objectives of this study were: 1) to evaluate and compare the necromass of “terra firme” and lowlands forests, (2) to study the relationship between necromass, above-ground biomass and wood density, and (3) to estimate the necromass of the department of Madre de Dios, Peru. Stocks of necromass and above-ground biomass were estimated at three different locations using permanent plots and line intercept transects. The average volume of necromass for the three sites was 72.9 m3 ha-1 with an average weight varying between 24.8 and 30.7 Mg ha-1, depending on the estimations of dead wood density used for the calculations. Terra firme forests had significantly higher stocks of necromass than lowland forests. The amount of necromass was 11% of the total above-ground biomass in Madre de Dios forests. The total stock of carbon stored in dead wood for the entire department of Madre de Dios was estimated to be approximately 100 mega tonnes of carbon. This is ten times more than the annual fossil fuel emissions of Peru between 2000 and 2008. The substantial stocks of necromass emphasize the importance of these types of field studies, considering that this component of tropical forest carbon cannot be detected using other methods such as satellite remote sensing.

Developing high-quality scientific research will be most effective if research communities with diverse skills and interests are able to share information and knowledge, are aware of the major challenges across disciplines, and can exploit economies of scale to provide robust answers and better inform policy. We evaluate opportunities and challenges facing the development of a more interactive research environment by developing an interdisciplinary synthesis of research on a single geographic region. We focus on the Amazon as it is of enormous regional and global environmental importance and faces a highly uncertain future. To take stock of existing knowledge and provide a framework for analysis we present a set of mini-reviews from fourteen different areas of research, encompassing taxonomy, biodiversity, biogeography, vegetation dynamics, landscape ecology, earth-atmosphere interactions, ecosystem processes, fire, deforestation dynamics, hydrology, hunting, conservation planning, livelihoods, and payments for ecosystem services. Each review highlights the current state of knowledge and identifies research priorities, including major challenges and opportunities. We show that while substantial progress is being made across many areas of scientific research, our understanding of specific issues is often dependent on knowledge from other disciplines. Accelerating the acquisition of reliable and contextualized knowledge about the fate of complex pristine and modified ecosystems is partly dependent on our ability to exploit economies of scale in shared resources and technical expertise, recognise and make explicit interconnections and feedbacks among sub-disciplines, increase the temporal and spatial scale of existing studies, and improve the dissemination of scientific findings to policy makers and society at large. Enhancing interaction among research efforts is vital if we are to make the most of limited funds and overcome the challenges posed by addressing large-scale interdisciplinary questions. Bringing together a diverse scientific community with a single geographic focus can help increase awareness of research questions both within and among disciplines, and reveal the opportunities that may exist for advancing acquisition of reliable knowledge. This approach could be useful for a variety of globally important scientific questions.

Lianas hold an important, but understudied, role in forest dynamics, however they are difficult to measure and detailed liana measurements are time consuming. Many researchers have therefore used an ordinal scale index, the crown occupancy index (COI), to describe the liana load carried by trees. Here we assess the overall effectiveness, in terms of accuracy, precision, repeatability and efficiency, of the COI in tropical forests. We relate the COI to more detailed liana measurements at the individual tree-level and site-level, comparing sites with different levels of liana infestation. Our results show (1) that the COI accurately measures individual tree and plot level liana loads, indicated by the strong correlations between the COI and the number and basal area of lianas. However, (2) as expected, the COI is only weakly related to the basal area of lianas rooted close to the tree, which is a proxy for competition for below-ground resources. The COI is also (3) an efficient measure of liana loads, as the input time needed for a COI survey is considerably less than that of a detailed liana survey. We also (4) found a high degree of repeatability in COI classification between observers. Additionally (5), the COI can be used to differentiate between sites in terms of their overall liana canopy competition (precision), but (6) may not be a precise indicator of the site-level mean basal area of lianas in tree crowns. © 2010 Elsevier B.V.

Photosynthetic leaf traits were determined for savanna and forest ecosystems in West Africa, spanning a large range in precipitation. Standardized major axis fits revealed important differences between our data and reported global relationships. Especially for sites in the drier areas, plants showed higher photosynthetic rates for a given N or P when compared with relationships from the global data set. The best multiple regression for the pooled data set estimated V(cmax) and J(max) from N(DW) and S. However, the best regression for different vegetation types varied, suggesting that the scaling of photosynthesis with leaf traits changed with vegetation types. A new model is presented representing independent constraints by N and P on photosynthesis, which can be evaluated with or without interactions with S. It assumes that limitation of photosynthesis will result from the least abundant nutrient, thereby being less sensitive to the allocation of the non-limiting nutrient to non-photosynthetic pools. The model predicts an optimum proportionality for N and P, which is distinct for V(cmax) and J(max) and inversely proportional to S. Initial tests showed the model to predict V(cmax) and J(max) successfully for other tropical forests characterized by a range of different foliar N and P concentrations.

*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.

Extensive areas of Amazonia undergo selective logging, modifying forest structure and nutrient cycles. Anthropogenic-accelerated rates of nitrogen (N) turnover could increase N loss and affect regeneration, carbon sequestration and timber production. We quantified leaf area reduction, canopy opening and downed biomass and resultant N flux from reduced impact logging (RIL) activities. We compared canopy reduction, surface soil moisture and nitrate to 8 m depth between logged gaps and intact primary forest to determine if logging activities increase subsoil nitrate. To test long-term logging effects, we evaluated surface N stocks along a 12-year postlogging chronosequence. At the harvest rate of 2.6 trees ha-1, total N additions in logging gaps, including leaves and wood from felled crowns (24.8 kg N ha-1) and other killed trees (41.9 kg N ha-1), accounted for over 80% of the total N addition to aboveground necromass from all logging activities (81.9 kg N ha-1). Despite this N turnover by logging, belowground nitrate storage to 8 m depth did not differ between logging gaps and primary forest at the low harvest rate and disturbance intensity of this study. Soil water depletion also did not differ between gaps and primary forest over 1 year, indicating the impact on belowground inorganic N was low. Compared with primary forest, nitrate concentrations to 8 m depth in logging gaps were only significantly higher at 60-100 cm, suggesting some N redistribution beyond the bulk of the fine roots in logging gaps. Extrapolated to the Amazon Basin scale, we provide a conservative estimate that logging damage and bole export under RIL would turn over 0.14 ± 0.07 to 0.23 ± 0.12 Tg N yr-1 based on 1999-2002 selective logging rates. Greater damage during conventional selective logging would cause higher N turnover throughout the Amazon Basin than our results based on RIL. © 2009 Blackwell Publishing Ltd.

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 response of terrestrial vegetation to a globally changing environment is central to predictions of future levels of atmospheric carbon dioxide. The role of tropical forests is critical because they are carbon-dense and highly productive. Inventory plots across Amazonia show that old-growth forests have increased in carbon storage over recent decades, but the response of one-third of the world's tropical forests in Africa is largely unknown owing to an absence of spatially extensive observation networks. Here we report data from a ten-country network of long-term monitoring plots in African tropical forests. We find that across 79 plots (163 ha) above-ground carbon storage in live trees increased by 0.63 Mg C ha(-1) yr(-1) between 1968 and 2007 (95% confidence interval (CI), 0.22-0.94; mean interval, 1987-96). Extrapolation to unmeasured forest components (live roots, small trees, necromass) and scaling to the continent implies a total increase in carbon storage in African tropical forest trees of 0.34 Pg C yr(-1) (CI, 0.15-0.43). These reported changes in carbon storage are similar to those reported for Amazonian forests per unit area, providing evidence that increasing carbon storage in old-growth forests is a pan-tropical phenomenon. Indeed, combining all standardized inventory data from this study and from tropical America and Asia together yields a comparable figure of 0.49 Mg C ha(-1) yr(-1) (n = 156; 562 ha; CI, 0.29-0.66; mean interval, 1987-97). This indicates a carbon sink of 1.3 Pg C yr(-1) (CI, 0.8-1.6) across all tropical forests during recent decades. Taxon-specific analyses of African inventory and other data suggest that widespread changes in resource availability, such as increasing atmospheric carbon dioxide concentrations, may be the cause of the increase in carbon stocks, as some theory and models predict.

A more detailed understanding of seasonal dynamics of carbon allocation between roots and shoots of temperate zone trees is needed. We labeled sapling stands of sugar maple (Acer saccharum Marsh) with 13CO2 in four 3 m diameter chambers during a 3 week interval in September 2006 and traced the movement of this 13C pulse through trees and soil during autumn and spring. High enrichment of 13C in foliage was achieved (δ13C = 387‰ ± 16‰). Late-season photosynthate was strongly mobilized during leaf senescence and stored in twigs, wood, and roots. Very high 13C enrichment of soil CO2 emissions at this time (δ13C = 766‰ ± 82‰) indicated the role of late-season photosynthesis in supplying root metabolism after the growing season. Rhizosphere soil was weakly enriched in 13C during fall, and increased significantly over the winter. Earthworms were highly enriched in fall, indicating that they consumed roots. In spring, 13C was strongly mobilized to growing shoots but not to growing fine roots; apparently, fine root growth in spring was not supplied by lateseason stored photosynthate. These results provide insights into the seasonal dynamics of temperate tree carbon allocation with implications for disruption of these dynamics by global warming.

Regional-scale above-ground biomass (AGB) estimates of tropical savannas and woodlands are highly uncertain, despite their global importance for ecosystems services and as carbon stores. In response, we collated field inventory data from 253 plots at four study sites in Cameroon, Uganda and Mozambique, and examined the relationships between field-measured AGB and cross-polarized radar backscatter values derived from ALOS PALSAR, an L-band satellite sensor. The relationships were highly significant, similar among sites, and displayed high prediction accuracies up to 150 Mg ha-1 (±∼20%). AGB predictions for any given site obtained using equations derived from data from only the other three sites generated only small increases in error. The results suggest that a widely applicable general relationship exists between AGB and L-band backscatter for lower-biomass tropical woody vegetation. This relationship allows regional-scale AGB estimation, required for example by planned REDD (Reducing Emissions from Deforestation and Degradation) schemes. Copyright 2009 by the American Geophysical Union.

This research represents one of the first studies in Amazonia to examine soil moisture and water-use efficiency (WUE) in secondary forest (SF) vegetation regrowing on abandoned pastures subjected to reduced nutrient constraints via a nutrient addition experiment. Extensive forested areas (about 80% of deforestation) have been converted to pastures in Amazonia, which were later abandoned following soil degradation and reduction in grass productivity. Colonization of these areas proceeds through species adapted to adverse edaphic conditions, such as low soil nutrients. Yet there is little data from such environments showing the interaction of soil nutrients and water availability on plant physiological processes. The objective of this study was to test whether three common SF tree species have positive physiological responses, e.g. increased photosynthesis and water-use efficiency, when nutrient limitations are relaxed through fertilization. The experiment was conducted on an abandoned pasture in central Amazonia with 6-year-old secondary vegetation following the application of four treatments: control; +phosphorus (P); +phosphorus and lime (P + Ca); and +phosphorus, lime and gypsum (P + Ca + G). The control had higher mean soil moisture at 140 and 180 cm depth at the end of the dry season, indicating that the treatment plots responded positively to fertilizer additions by taking up additional water. Trees of Vismia japurensis and Bellucia grossularioides growing on the fertilized plots had the highest net photosynthesis rates (A) (18.7 and 20.4 μmol m-2 s-1, respectively). The three species utilized different strategies with regard to physiological and nutritional response, with V. japurensis, regardless of treatment, using these limiting resources most efficiently to colonize abandoned pastures. Trees growing on the P + Ca but not +P alone plots increased a rates, indicating that Ca is an important limiting nutrient in post-pasture secondary succession. The addition of Ca as ash by burning primary and secondary vegetation could explain the rapid growth and dominance of V. japurensis in abandoned pasture areas in central Amazonia. The efficiency of Vismia to use limiting resources could lead to a restructuring of SF and altered rates of stand-level productivity. © 2008 Elsevier B.V. All rights reserved.

Question: is soil hydrology an important niche-based driver of biodiversity in tropical forests? More specifically, we asked whether seasonal dynamics in soil water regime contributed to vegetation partitioning into distinct forest types. Location: Tropical rain forest in northwestern Mato Grosso, Brazil. Methods: We investigated the distribution of trees and lianas ≥ 1 cm DBH in ten transects that crossed distinct hydrological transitions. Soil water content and depth to water table were measured regularly over a 13-month period. Results: a detrended correspondence analysis (DCA) of 20 dominant species and structural attributes in 10 x 10 m subplots segregated three major forest types: (1) high-statured upland forest with intermediate stem density, (2) medium-statured forest dominated by palms, and (3) low-statured campinarana forest with high stem density. During the rainy season and transition into the dry season, distinct characteristics of the soil water regime (i.e. hydro-indicators) were closely associated with each vegetation community. Stand structural attributes and hydro-indicators were statistically different among forest types. Conclusions: Some upland species appeared intolerant of anaerobic conditions as they were not present in palm and campinarana sites, which experienced prolonged periods of saturation at the soil surface. A shallow impermeable layer restricted rooting depth in the campinarana community, which could heighten drought stress during the dry season. The only vegetation able to persist in campinarana sites were short-statured trees that appear to be well-adapted to the dual extremes of inundation and drought. © IAVS; Opulus Press.

Nearly all published rates of secondary forest (SF) regrowth for Amazonia are inferred from chronosequences. We examined SF regrowth on abandoned pastures over a 4-year period to determine if measured rates of forest recovery differ from chronosequence predictions. We studied the emergence, development and death of over 1300 stems in 10 SFs representing three age classes (

The purpose of this study was to determine if spatially-explicit commercial timber inventories (CTI) could be used in conjunction with satellite imagery to improve timber assessments and forest biomass estimates in Amazonia. As part of a CTI, all commercial trees ≥45 cm DBH were measured and georeferenced in 3500 ha of a logging concession in NW Mato Grosso, Brazil. A scientific inventory was conducted of all trees and palms ≥10 cm DBH in 11.1 ha of this area. A total of >20,000 trees were sampled for both inventories. To characterize vegetation radiance and topographic features, regional LANDSAT TM and ASTER images were obtained. Using a stream network derived from the ASTER-based 30 m digital elevation model (DEM), a procedure was developed to predict areas excluded from logging based on reduced impact logging (RIL) criteria. A topographic index (TI) computed from the DEM was used to identify areas with similar hydrologic regimes and to distinguish upland and lowland areas. Some timber species were associated with convergent landscape positions (i.e. higher TI values). There were significant differences in timber density and aboveground biomass (AGB) in upland (6.0 stems ha-1, 33 Mg ha-1) versus lowland (5.4 stems ha-1, 29 Mg ha-1) areas. Upland and lowland, and timber and non-timber areas could be distinguished through single and principal component analysis of LANDSAT bands. However, radiance differences between areas with and without commercial timber on a sub-hectare scale were small, indicating LANDSAT images would have limited utility for assessing commercial timber distribution at this scale. Assuming a 50 m stream buffer, areas protected from logging ranged from 7% (third order streams and above) to 28% (first order and above) of the total area. There was a strong positive relationship between AGB based on the scientific inventory of all trees and from the commercial timber, indicating that the CTI could be used in conjunction with limited additional sampling to predict total AGB (276 Mg ha-1). The methods developed in this study could be useful for facilitating commercial inventory practices, understanding the relationship of tree species distribution to landscape features, and improving the novel use of CTIs to estimate AGB. © 2006 Elsevier B.V. All rights reserved.

The success of initial tree seedling establishment is related to the capture and use of primary resources such as light and nutrients. The selection of tree species with a greater potential to assimilate carbon and capacity to efficiently utilize nutrients and light would facilitate the revegetation of degraded areas, primarily where irradiance is high and soil nutrient availability low. We analyzed soil physical and chemical characteristics, survival, growth, photosynthesis, chlorophyll a fluorescence, leaf macro- and micro-nutrient content and photosynthetic nutrient use efficiency in young tropical tree species planted in degraded areas in central Amazonia. The species studied were: Bellucia grossularioides, Bombacopsis macrocalyx, Cecropia ficifolia, Cecropia sciadophylla, Chrysophyllum sanguinolentum, Eugenia cumini, Inga edulis and Iryanthera macrophyla. C. sanguinolentum, a late secondary species, exhibited the lowest survival rates, except when compared with B. grossularioides and C. sciadophylla. Photosynthesis varied between 34 and 264 nmol g-1 s-1 for the eight species; species of Cecropia had photosynthetic values eight times greater than C. sanguinolentum. For the photochemical efficiency of photosystem II, C. sanguinolentum presented the highest degree of photoinhibition as a result of an inefficient use of excess irradiance. Leaf macronutrient concentrations varied from 16 to 29, 0.4 to 1.0, 6 to 13, 7 to 22, 1.6 to 3.4 g kg-1 for N, P, K, Ca and Mg, respectively. For photosynthetic nutrient use efficiency, in general, the two species of Cecropia exhibited the highest values, while C. sanguinolentum presented the lowest macronutrient use efficiency. The two species of Cecropia had photosynthetic nitrogen use efficiency seven times greater than C. sanguinolentum. On these highly degraded soils we found that pioneer trees (Cecropia) are better able to colonize than species from other successional stages. Trees on these degraded soils are primarily limited by P or micronutrients. Despite the remove of the O horizon, N does not appear to limit photosynthetic activity. From this we conclude that species of Cecropia possess ecophysiological mechanisms associated with carbon assimilation and nutrient use that determines success in early establishment and has potential to recuperate degraded areas, especially when compared to C. sanguinolentum. Early secondary species such as I. edulis and E. cumini could also be used with success in forest plantings to recuperate degraded areas when selecting for species, (1) efficient in the utilization of excess energy for photosynthesis, (2) efficient in the use of limited soil nutrients and (3) with high survival and growth rates. © 2006 Elsevier B.V. All rights reserved.

Large areas of Amazonian forest have been converted to pastures over the last two decades. Low soil fertility and mismanagement results in a rapid decline in net primary productivity leading the pastures to be abandoned, and woody vegetation adapted to low nutrient conditions colonize the areas. The objective of this study was to examine nutrient use efficiency, following liming (calcium) and phosphorus addition by three of the most frequent colonizing species. The experiment conducted on a six year-old secondary forest, consisted of four treatments: control; phosphorus addition (P); phosphorus and lime addition (P+Cal); and phosphorus, lime and gypsum addition (P+Cal+G). Leaf gas exchange, soil and leaf nutrient concentration were determined eight months after the treatment application. There was a significant response by species to the addition of phosphorus and lime (P+Cal and P+Cal+G). The species, Bellucia grossularioides accumulated more N, P and Zn in the leaves, while Laetia procera accumulated more Ca and Mn. The species Vismia japurensis had higher nutrient use efficiency, as a function of the higher photosynthetic rates. Vismia japurensis presented lower P concentrations than Bellucia grossularioides, suggesting that is well adapted to environments low in nutrients, as this species often occurs in degraded areas in Amazonia.

The area of secondary forest (SF) regenerating from pastures is increasing in the Amazon basin; however, the return of forest and canopy structure following abandonment is not well understood. This study examined the development of leaf area index (LAI), canopy cover, aboveground biomass, stem density, diameter at breast height (DBH), and basal area (BA) by growth form and diameter class for 10 SFs regenerating from abandoned pastures. Biomass accrual was tree dominated, constituting ≥94% of the total measured biomass in all forests abandoned ≥4 to 6 yr. Vine biomass increased with forest age, but its relative contribution to total biomass decreased with time. The forests were dominated by the tree Vismia spp. (>50%). Tree stem density peaked after 6 to 8 yr (10 320 stems per hectare) before declining by 42% in the 12- to 14-yr-old SFs. Small-diameter tree stems in the 1-5-cm size class composed >58% of the total stems for all forests. After 12 to 14 yr, there was no significant leaf area below 150-cm height. Leaf area return (LAI = 3.2 after 12 to 14 yr) relative to biomass was slower than literature-reported recovery following slash-and-burn, where LAI can reach primary forest levels (LAI = 4-6) in 5 yr. After 12 to 14 yr, the colonizing vegetation returned some components of forest structure to values reported for primary forest. Basal area and LAI were 50%-60%, canopy cover and stem density were nearly 100%, and the rapid tree-dominated biomass accrual was 25%-50% of values reported for primary forest. Biomass accumulation may reach an asymptote earlier than expected because of even-aged, monospecific, untiered stand structure. The very slow leaf area accumulation relative to biomass and to reported values for recovery following slash-and-burn indicates a different canopy development pathway that warrants further investigation of causes (e.g. nutrient limitations, competition) and effects on processes such as evapotranspiration and soil water uptake, which would influence long-term recovery rates and have regional implications.

Amazonian farmers and ranchers use fire to clear land for agriculture and pasture as part of extensive land-use strategies that have deforested 500 000 km2 over the past 25 years. Ash from burning biomass fertilizes crops and pastures, but declining productivity often occurs after a few years, generally leading to land abandonment and further clearing. Subsequent forest regrowth partially offsets carbon emissions from deforestation, but is often repeatedly cleared and burned. In the first quantitative, basin-wide assessment of the effect of repeated clearing and burning on forest regrowth, our analysis of data from 90 stands at nine locations across the region indicates that stands with a history of five or more fires suffer on average a greater than 50% reduction in carbon accumulation. In the absence of management interventions, Amazonian landscapes dominated by this pronounced legacy of fire are apt to accumulate very little carbon and will remain highly susceptible to recurrent burning. © the Ecological Society of America.

We examined carbon export in whole logs and carbon accumulation as coarse woody debris (CWD) produced from forest damage during all phases of the first and second year of a certified reduced impact logging (RIL) timber harvest in southern Amazonia. Our measurements included a 100% survey of roads and log decks, assessment of canopy damage and ground disturbance in skid trails and tree-fall gaps, and measurement of carbon exported from the site in logs. Log deck and road construction crushed one and five trees in the 10-60 cm diameter at breast height (DBH) class per hectare logged, disturbed areas of 24 and 100 m2 ha-1, respectively, and together disturbed about 1% of the forest. On average 1.1-2.6 trees ha-1 were harvested over the two years. Logged gaps constituted the greatest disturbance on an area basis (4-10% of the forest) and CWD generation (1.9-4.4 Mg ha-1 logged). In gaps, felled trees severed or crushed 10 trees ≥10 cm DBH per tree logged, which corresponded to 1.7 Mg ha-1 of CWD per tree logged. Crown height - measured from the first bifurcation to the top of the crown - rather than tree height was the better predictor of gap size formed from tree felling (R 2 = 0.41). Logging activities significantly reduced leaf area in roads, log decks and gaps, with the greatest reduction (48%) in log decks and least in logged gaps and roads (28-33%) compared to undisturbed forest. A total of 37 species were harvested, with 36% of the total trees harvested and 48% of the total carbon exported from the site in three of the most common species. Logging damage produced 4.9-8.8 Mg C ha-1 logged of CWD from all phases of the operation. Carbon export in whole logs (2.1-3.7 Mg C ha -1 logged) represented 1-3% of the total standing forest carbon ≥10 cm DBH (138 Mg C ha-1). The mean carbon ratio (per hectare logged) of C in CWD to C exported in logs was 2.4. The disturbance, damage, carbon export and CWD data we present advances understanding of the effect of selective logging on tropical forest dynamics of the Amazon Basin. Our results indicate that certified timber harvest in Amazonia under RIL is a viable forest management option to reduce damage and CWD production compared to conventional logging (CL) practices; however, the benefits of disturbance reduction from RIL relative to CL are only realized at greater volumes of timber extraction. © 2005 Elsevier B.V. All rights reserved.

Over the past three decades, large expanses of forest in the Amazon Basin were converted to pasture, many of which later degraded to woody fallows and were abandoned. While the majority of tropical secondary forest (SF) studies have examined post-deforestation or post-agricultural succession, we examined post-pasture forest recovery in 10 forests ranging in age from 0 to 14 years since abandonment. We measured aboveground biomass and soil nutrients to 45 cm depth and computed total site carbon (C) and nutrient stocks to gain an understanding of the dynamics of nutrient and C buildup in regenerating SF in central Amazonia. Aboveground biomass accrual was rapid, 11.0 Mg·ha -1·yr-1, in the young SFs. Within 12-14 yr, they accumulated up to 128.1 Mg/ha of dry aboveground biomass, equivalent to 25-50% of primary forest biomass in the region. Wood nitrogen (N) and phosphorus (P) concentrations decreased with forest age. Aboveground P and calcium (Ca) stocks accumulated at a rate of 1.2 and 29.4 kg·ha-1·yr -1; extractable soil P stocks declined as forest age increased. Although soil stocks of exchangeable Ca (207.0 ± 23.7 kg/ha) and extractable P (8.3 ± 1.5 kg/ha) were low in the first 45 cm, both were rapidly translocated from soil to plant pools. Soil N stocks increased with forest age, probably due to N fixation, atmospheric deposition, and/or subsoil mining. Total soil C storage to 45 cm depth ranged between 42 and 84 Mg/ha, with the first 15 cm storing 40-45% of the total. Total C accrual (7.04 Mg C·ha-1·yr-1) in both aboveground and soil pools was similar or higher than values reported in other studies. Tropical SFs regrowing on lightly to moderately used pasture rapidly sequester C and rebuild total nutrient capital following pasture abandonment. Translocation of some nutrients from deep soil (>45 cm depth) may be important to sustaining productivity and continuing biomass accumulation in these forests. The soil pool represents the greatest potential for long-term C gains; however, soil nutrient deficits may limit future productivity.

The Large-scale Biosphere-Atmosphere Experiment in Amazonia (LBA) is a multinational, interdisciplinary research program led by Brazil. Ecological studies in LBA focus on how tropical forest conversion, regrowth, and selective logging influence carbon storage, nutrient dynamics, trace gas fluxes, and the prospect for sustainable land use in the Amazon region. Early results from ecological studies within LBA emphasize the variability within the vast Amazon region and the profound effects that land-use and land-cover changes are having on that landscape. The predominant land cover of the Amazon region is evergreen forest; nonetheless, LBA studies have observed strong seasonal patterns in gross primary production, ecosystem respiration, and net ecosystem exchange, as well as phenology and tree growth. The seasonal patterns vary spatially and interannually and evidence suggests that these patterns are driven not only by variations in weather but also by innate biological rhythms of the forest species. Rapid rates of deforestation have marked the forests of the Amazon region over the past three decades. Evidence from ground-based surveys and remote sensing show that substantial areas of forest are being degraded by logging activities and through the collapse of forest edges. Because forest edges and logged forests are susceptible to fire, positive feedback cycles of forest degradation may be initiated by land-use-change events. LBA studies indicate that cleared lands in the Amazon, once released from cultivation or pasture usage, regenerate biomass rapidly. However, the pace of biomass accumulation is dependent upon past land use and the depletion of nutrients by unsustainable land-management practices. The challenge for ongoing research within LBA is to integrate the recognition of diverse patterns and processes into general models for prediction of regional ecosystem function.