Accurate characterization of the role of the dry tropics in the global carbon cycle requires precise estimation of woody biomass changes due to ecological and anthropogenic change, including deforestation, forest degradation, regrowth, mortality and enhanced tree growth due to climate change. L-band Synthetic Aperture Radar (SAR) backscatter observations offer a reliable option to consistently map these processes as they are (i) available globally since 2007 (JAXA ALOS-1, ALOS-2 and ALOS-4), and (ii) sensitive to woody structure, such as aboveground biomass density (AGBD) up to similar to 100 t ha(-1). However, we lack multi-site empirical understanding of the scattering processes that determine the relationship between L-band SAR and woody vegetation structure in the dry tropics, and how this is mediated by soil properties. This study used observations from ground plots in Africa (n = 171), Australia (n = 6), and South America (n = 44) to understand the impact of vegetation structure and soil properties on spatially and temporally coincident fully-polarimetric L-band SAR data. Fully-polarimetric L-band SAR single-look complex data were converted to scattering mechanisms/parameters using van Zyl, Cloude-Pottier, and Freeman-Durden polarimetric decompositions to elucidate the physical mechanisms involved. Multivariate SAR-vegetation-soil relationships were analysed using a theory-informed structural equation modelling approach. The strongest positive effects on volume scattering come from stem density (stems ha(-1)) and mean stem biomass of trees, and soil water and sand content (standardized regression coefficients of 0.3, 0.1, 0.2 and 0.1, respectively). The only significant effect on surface scattering is from stem density (0.1). Significant effects on double bounce scattering are from stem density (0.3) and soil sand content (-0.2). Since AGBD is the product of stem density and mean stem biomass, this modelling framework points to a stronger effect from the number of trees rather than their size/biomass. Therefore, AGBD maps relying solely on radar intensity may not reflect significant changes when AGBD is increasing due to the growth of existing stems. Additionally, such maps might overestimate changes in AGBD when driven by the recruitment of new stems or loss of existing stems. Full-polarimetric observations allow the decomposition of the radar signal into volume scattering, surface scattering, and double bounce, enabling the inversion of structural equation models to retrieve both stem density and mean stem biomass. This provides a more comprehensive description of forest structure compared to retrieving only AGBD. As this approach depends on full-polarimetric data, its effectiveness is closely tied to the availability of such observations. Our findings underscore the value of recent and upcoming missions such as ALOS-4 PALSAR-3, BIOMASS and ROSE-L, and highlight the need to prioritise the acquisition of quad-pol SAR data to support future large-scale retrieval of vegetation structure attributes.
ABSTRACT Aim We investigate the distribution of dispersal modes — endozoochory (via animal ingestion), synzoochory (via animal seed accumulation) and anemochory (via wind)—and seed mass of woody plants across Brazilian ecosystems, testing whether their relative abundance in local plant communities could be predicted by climate, edaphic factors and dispersal agents (e.g., the main vectors moving seeds). We expected climate to be the main predictor of dispersal strategy abundance. Location Brazilian ecosystems (Amazon, Atlantic Forest, Cerrado and Caatinga). Time Period 2004–2020. Major Taxa Studied Woody angiosperms, trees and shrubs. Methods We assigned seed mass to 1984 species and dispersal mode to 2203 species within 300 old‐growth and non‐flooded plots. We employed Random Forest regressions to assess the influence of water availability, temperature seasonality, soil fertility, wind and frugivorous primates on the community‐weighted means of seed mass and dispersal modes. We mapped predictions and estimated the area of applicability to identify regions where predictions were reliable. Results Climate was the main factor predicting tree dispersal strategies, followed by dispersal agents or soil properties. Endozoochory was prevalent in wetter regions with high water availability and greater diurnal temperature range. Anemochory was more common in drier regions with low water availability, stronger winds and greater diurnal temperature range, whereas synzoochory was more abundant in areas with low temperature seasonality. Heavier seeds were more prevalent in regions with less seasonal temperatures, smaller diurnal temperature ranges and weaker winds. Main Conclusions Climate is a key predictor of the distribution and prevalence of plants with contrasting dispersal modes and seed masses. We thus expect functional responses to environmental change associated with shifts in the relative abundance of dispersal strategies, with implications for ecosystem services. Our results also highlight the critical role of plot‐based datasets in improving our understanding of how environmental factors influence seed dispersal strategies across tropical ecosystems.
Tree growth and longevity trade-offs fundamentally shape the terrestrial carbon balance. Yet, we lack a unified understanding of how such trade-offs vary across the world’s forests. By mapping life history traits for a wide range of species across the Americas, we reveal considerable variation in life expectancies from 10 centimeters in diameter (ranging from 1.3 to 3195 years) and show that the pace of life for trees can be accurately classified into four demographic functional types. We found emergent patterns in the strength of trade-offs between growth and longevity across a temperature gradient. Furthermore, we show that the diversity of life history traits varies predictably across forest biomes, giving rise to a positive relationship between trait diversity and productivity. Our pan-latitudinal assessment provides new insights into the demographic mechanisms that govern the carbon turnover rate across forest biomes.
Using 2.046 botanically-inventoried tree plots across the largest tropical forest on Earth, we mapped tree species-diversity and tree species-richness at 0.1-degree resolution, and investigated drivers for diversity and richness. Using only location, stratified by forest type, as predictor, our spatial model, to the best of our knowledge, provides the most accurate map of tree diversity in Amazonia to date, explaining approximately 70% of the tree diversity and species-richness. Large soil-forest combinations determine a significant percentage of the variation in tree species-richness and tree alpha-diversity in Amazonian forest-plots. We suggest that the size and fragmentation of these systems drive their large-scale diversity patterns and hence local diversity. A model not using location but cumulative water deficit, tree density, and temperature seasonality explains 47% of the tree species-richness in the terra-firme forest in Amazonia. Over large areas across Amazonia, residuals of this relationship are small and poorly spatially structured, suggesting that much of the residual variation may be local. The Guyana Shield area has consistently negative residuals, showing that this area has lower tree species-richness than expected by our models. We provide extensive plot meta-data, including tree density, tree alpha-diversity and tree species-richness results and gridded maps at 0.1-degree resolution.
The tropical forest carbon sink is known to be drought sensitive, but it is unclear which forests are the most vulnerable to extreme events. Forests with hotter and drier baseline conditions may be protected by prior adaptation, or more vulnerable because they operate closer to physiological limits. Here we report that forests in drier South American climates experienced the greatest impacts of the 2015–2016 El Niño, indicating greater vulnerability to extreme temperatures and drought. The long-term, ground-measured tree-by-tree responses of 123 forest plots across tropical South America show that the biomass carbon sink ceased during the event with carbon balance becoming indistinguishable from zero (−0.02 ± 0.37 Mg C ha −1 per year). However, intact tropical South American forests overall were no more sensitive to the extreme 2015–2016 El Niño than to previous less intense events, remaining a key defence against climate change as long as they are protected.
In a time of rapid global change, the question of what determines patterns in species abundance distribution remains a priority for understanding the complex dynamics of ecosystems. The constrained maximization of information entropy provides a framework for the understanding of such complex systems dynamics by a quantitative analysis of important constraints via predictions using least biased probability distributions. We apply it to over two thousand hectares of Amazonian tree inventories across seven forest types and thirteen functional traits, representing major global axes of plant strategies. Results show that constraints formed by regional relative abundances of genera explain eight times more of local relative abundances than constraints based on directional selection for specific functional traits, although the latter does show clear signals of environmental dependency. These results provide a quantitative insight by inference from large-scale data using cross-disciplinary methods, furthering our understanding of ecological dynamics.
For more than three decades, major efforts in sampling and analyzing tree diversity in South America have focused almost exclusively on trees with stems of at least 10 and 2.5 cm diameter, showing highest species diversity in the wetter western and northern Amazon forests. By contrast, little attention has been paid to patterns and drivers of diversity in the largest canopy and emergent trees, which is surprising given these have dominant ecological functions. Here, we use a machine learning approach to quantify the importance of environmental factors and apply it to generate spatial predictions of the species diversity of all trees (dbh ≥ 10 cm) and for very large trees (dbh ≥ 70 cm) using data from 243 forest plots (108,450 trees and 2832 species) distributed across different forest types and biogeographic regions of the Brazilian Amazon. The diversity of large trees and of all trees was significantly associated with three environmental factors, but in contrasting ways across regions and forest types. Environmental variables associated with disturbances, for example, the lightning flash rate and wind speed, as well as the fraction of photosynthetically active radiation, tend to govern the diversity of large trees. Upland rainforests in the Guiana Shield and Roraima regions had a high diversity of large trees. By contrast, variables associated with resources tend to govern tree diversity in general. Places such as the province of Imeri and the northern portion of the province of Madeira stand out for their high diversity of species in general. Climatic and topographic stability and functional adaptation mechanisms promote ideal conditions for species diversity. Finally, we mapped general patterns of tree species diversity in the Brazilian Amazon, which differ substantially depending on size class.
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 <5 m) had 18% less above-ground woody productivity and 23% less biomass stock than forests with a deep water table. Forests in drier climates (maximum cumulative water deficit < -160 mm) had 21% less productivity and 24% less biomass than those in wetter climates. Productivity was affected by the interaction between climatic water deficit and water table depth. On average, in drier climates the forests with a shallow water table had lower productivity than those with a deep water table, with this difference decreasing within wet climates, where lower productivity was confined to a very shallow water table. Main conclusions We show that the two extremes of water availability (excess and deficit) both reduce productivity in Amazon upland (terra-firme) forests. Biomass and productivity across Amazonia respond not simply to regional climate, but rather to its interaction with water table conditions, exhibiting high local differentiation. Our study disentangles the relative contribution of those factors, helping to improve understanding of the functioning of tropical ecosystems and how they are likely to respond to climate change.
The 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.
In a time of rapid global change, the question of what determines patterns in species abundance distribution remains a priority for understanding the complex dynamics of ecosystems. The constrained maximization of information entropy provides a framework for the understanding of such complex systems dynamics by a quantitative analysis of important constraints via predictions using least biased probability distributions. We apply it to over two thousand hectares of Amazonian tree inventories across seven forest types and thirteen functional traits, representing major global axes of plant strategies. Results show that constraints formed by regional relative abundances of genera explain almost ten times more of local relative abundances then constraints based on either directional or stabilizing selection for specific functional traits, although the latter does show clear signals of environmental dependency. These results provide a quantitative insight by inference from large-scale data using cross-disciplinary methods, furthering our understanding of ecological dynamics.
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.
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.
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.
ABSTRACT: 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 rate of decline 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.
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.
There is a consensus about negative impacts of droughts in Amazonia. Yet, extreme wet episodes, which are becoming as severe and frequent as droughts, are overlooked and their impacts remain poorly understood. Moreover, drought reports are mostly based on forests over a deep water table (DWT), which may be particularly sensitive to dry conditions. Based on demographic responses of 30 abundant tree species over the past two decades, in this study we analyzed the impacts of severe droughts but also of concurrent extreme wet periods, and how topographic affiliation (to shallow - SWTs - or deep - DWTs - water tables), together with species functional traits, mediated climate effects on trees. Dry and wet extremes decreased growth and increased tree mortality, but interactions of these climatic anomalies had the highest and most positive impact, mitigating the simple negative effects. Despite being more drought-tolerant, species in DWT forests were more negatively affected than hydraulically vulnerable species in SWT forests. Interaction of wet-dry extremes and SWT depth modulated tree responses to climate, providing buffers to droughts in Amazonia. As extreme wet periods are projected to increase and at least 36% of the Amazon comprises SWT forests, our results highlight the importance of considering these factors in order to improve our knowledge about forest resilience to climate change.
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.
EVI in water-limited sites + pre R² = 0.61 + cld R² = 0.48 -dtr R² = 0.62 + vap R² = 0.55 + tmn R² = 0.42 + swc R² = 0.54 + rad R² = 0.31 ± pet R² = 0.36 ± tmp R² = 0.24 ± tmx R² = 0.23 a EVI in sites with mixed limitations + pre R² = 0.13 + cld R² = 0.08 ± dtr R² = 0.09 + vap R² = 0.18 + tmn R² = 0.22 ± swc R² = 0.17 ± rad R² = 0.33 ± pet R² = 0.14 ± tmp R² = 0.18 ± tmx R² = 0.13
Abstract. The seasonal climate drivers of the carbon cycle in tropical forests remain poorly known, although these forests account for more carbon assimilation and storage than any other terrestrial ecosystem. Based on a unique combination of seasonal pan-tropical data sets from 89 experimental sites (68 include aboveground wood productivity measurements and 35 litter productivity measurements), their associate canopy photosynthetic capacity (enhanced vegetation index, EVI) and climate, we ask how carbon assimilation and aboveground allocation are related to climate seasonality in tropical forests and how they interact in the seasonal carbon cycle. We found that canopy photosynthetic capacity seasonality responds positively to precipitation when rainfall is < 2000 mm.yr−1 (water-limited forests) and to radiation otherwise (light-limited forests); on the other hand, independent of climate limitations, wood productivity and litterfall are driven by seasonal variation in precipitation and evapotranspiration respectively. Consequently, light-limited forests present an asynchronism between canopy photosynthetic capacity and wood productivity. Precipitation first-order control indicates an overall decrease in tropical forest productivity in a drier climate.
The seasonal climate drivers of the carbon cycle in tropical forests remain poorly known, although these forests account for more carbon assimilation and storage than any other terrestrial ecosystem. Based on a unique combination of seasonal pan-tropical data sets from 89 experimental sites (68 include aboveground wood productivity measurements and 35 litter productivity measurements), their associated canopy photosynthetic capacity (enhanced vegetation index, EVI) and climate, we ask how carbon assimilation and aboveground allocation are related to climate seasonality in tropical forests and how they interact in the seasonal carbon cycle. We found that canopy photosynthetic capacity seasonality responds positively to precipitation when rainfall is < 2000 mm yr(-1) (water-limited forests) and to radiation otherwise (light-limited forests). On the other hand, independent of climate limitations, wood productivity and litterfall are driven by seasonal variation in precipitation and evapotranspiration, respectively. Consequently, light-limited forests present an asynchronism between canopy photosynthetic capacity and wood productivity. First-order control by precipitation likely indicates a decrease in tropical forest productivity in a drier climate in water-limited forest, and in current light-limited forest with future rainfall < 2000 mm yr(-1).