Old-growth tropical forests store vast amounts of carbon in their aboveground biomass (AGB), yet the relative roles of abiotic factors such as climate, soil, and topography in governing its spatial distribution remain poorly understood. In particular, the degree to which climate acts on AGB through forest structure is still poorly quantified at the pantropical scale. Using a pantropical dataset of more than 2,000 old-growth forest plots and a structure-explicit framework, we assess how climate influences AGB through its effects on four structural attributes: basal area, mean diameter, stem density, and basal area-weighted wood density. We find that climate shapes AGB primarily through its effects on forest structure. However, structural attributes respond to climate in opposite directions, so climate’s net effect on AGB largely cancels out, and no clear climate-AGB relationship emerges across tropical regions. Moreover, only wood density responds consistently, decreasing with annual precipitation and increasing with precipitation seasonality, whereas all other attributes respond to climate differently from one region to another. This geographical variation further obscures any global climatic signal on AGB and points to the role of biogeographic history in shaping forest structure. Our findings highlight the central role of the climate-structure nexus in explaining AGB variation, and call for structure-explicit models to improve carbon stock predictions and inform climate adaptation strategies.

Amazon rainforests face intensifying water stress due to increases in vapour pressure deficit and changing hydrological regimes. Embolism resistance (Ψ50) is a critical metric of tree survival under drought conditions, it is defined as a plant's capacity to resist disruption of xylem water flow due to air bubble formation from water stress. However, measurements of Ψ50 are only available for a limited number of Amazon locations and species. Conversely, data on forest taxonomic composition are abundant across Amazonia, and if Ψ50 is conserved phylogenetically, these data could provide a way to scale-up drought resistance patterns. Here we evaluate Ψ50 measurements across non-flooded Amazonian tree taxa and reveal a moderate phylogenetic signal, with phylogenetic conservatism evident at the family-level. Notably, Fabaceae is amongst the most embolism-resistant tree families in Amazonia. Leveraging the phylogenetic signal we use species composition and tree size data from 448 forest plots across Amazonia to produce a macroecological assessment of Amazonian vulnerability to embolism. The resulting estimate spatial pattern reveals that forests in the Brazilian and Guiana Shield regions, where Fabaceae abundance is high, show strong resistance to embolism. In contrast, tree communities in Western Amazonia appear more vulnerable to embolism, suggesting a reduced capacity to withstand future drought conditions.

Wood density is a critical control on tree biomass, so poor understanding of its spatial variation can lead to large and systematic errors in forest biomass estimates and carbon maps. The need to understand how and why wood density varies is especially critical in tropical America where forests have exceptional species diversity and spatial turnover in composition. As tree identity and forest composition are challenging to estimate remotely, ground surveys are essential to know the wood density of trees, whether measured directly or inferred from their identity. Here, we assemble an extensive dataset of variation in wood density across the most forested and tree-diverse continent, examine how it relates to spatial and environmental variables, and use these relationships to predict spatial variation in wood density over tropical and sub-tropical South America. Our analysis refines previously identified east-west Amazon gradients in wood density, improves them by revealing fine-scale variation, and extends predictions into Andean, dry, and Atlantic forests. The results halve biomass prediction errors compared to a naïve scenario with no knowledge of spatial variation in wood density. Our findings will help improve remote sensing-based estimates of aboveground biomass carbon stocks across tropical South America.

Understanding how the traits of lineages are related to diversification is key for elucidating the origin of variation in species richness. Here, we test whether traits are related to species richness among lineages of trees from all major biogeographical settings of the lowland wet tropics. We explore whether variation in mortality rate, breeding system and maximum diameter are related to species richness, either directly or via associations with range size, among 463 genera that contain wet tropical forest trees. For Amazonian genera, we also explore whether traits are related to species richness via variation among genera in mean species-level range size. Lineages with higher mortality rates—faster life-history strategies—have larger ranges in all biogeographic settings and have higher mean species-level range sizes in Amazonia. These lineages also have smaller maximum diameters and, in the Americas, contain dioecious species. In turn, lineages with greater overall range size have higher species richness. Our results show that fast life-history strategies influence species richness in all biogeographic settings because lineages with these ecological strategies have greater range sizes. These links suggest that dispersal has been a key process in the evolution of the tropical forest flora.

Understanding how functional traits are related to species diversity and ecosystem properties is a central goal of ecology. Wood density is a trait that integrates many aspects of plant form and function and is highly variable among species. Previous studies of wood density across elevational gradients have been based on limited sampling and have reported declines with increasing elevation, though even this simple pattern remains unknown, much less its underlying functional and evolutionary relationships. Here, we use one of the longest and most speciose elevational gradients in the world, extending from the Andean tree line to the Amazon basin, to test the extent to which elevation, species composition, phylogenetic affinity and forest structure determine variation in wood density. Using field-collected wood samples and global databases, we assigned wood density to 1231 species and 31,330 stems across 41 (47.5 ha) mature forest plots arrayed across a 3500-m vertical gradient. Our results show that mean wood density, either weighted by abundance, basal area or species, was highly variable but tended to decline from low to middle elevations and increase again from mid-elevations to the tree line. As a result of this non-linearity, forests at the Andean tree line had higher wood density than their lowland Amazon counterparts. We observed an abrupt transition in wood density at the lower limit of persistent cloud formation (cloud base), where the lowest wood density values were found. The decline of wood density is attributed to a significant shift in life forms, with an abundance of tree ferns at middle elevations and a higher probability of landslides and disturbances favouring a suite of traits associated with low wood density, such as softer wood and higher elasticity. Species turnover explained most of the among-species variation across the gradient, with elevation having no consistent effect on within-species variation in wood density. Together, both gradual compositional changes and sharp local changes in the importance of non-dicot life forms, such as arborescent ferns and palms, define patterns of forest-level carbon density, with wood density per se controlling ecosystem properties, such as carbon flux, across the Andes-to-Amazon elevational gradient.

Climate change is shifting species distributions, leading to changes in community composition and novel species assemblages worldwide. However, the responses of tropical forests to climate change across large-scale environmental gradients remain largely unexplored. Using long-term data over 66,000 trees of more than 2,500 species occurring over 3,500 m elevation along the hyperdiverse Amazon-to-Andes elevational gradients in Peru and Bolivia, we assessed community-level shifts in species composition over a 40+ y time span. We tested the thermophilization hypothesis, which predicts an increase in the relative abundances of species from warmer climates through time. Additionally, we examined the relative contributions of tree mortality, recruitment, and growth to the observed compositional changes. Mean thermophilization rates (TR) across the Amazon-to-Andes gradient were slow relative to regional temperature change. TR were positive and more variable among Andean forest plots compared to Amazonian plots but were highest at midelevations around the cloud base. Across all elevations, TR were driven primarily by tree mortality and decreased growth of highland (cool-adapted) species rather than an influx of lowland species with higher thermal optima. Given the high variability of community-level responses to warming along the elevational gradients, the high tree mortality, and the slower-than-warming rates of compositional change, we conclude that most tropical tree species, and especially lowland Amazonian tree species, will not be able to escape current or future climate change through upward range shifts, causing fundamental changes to composition and function in Earth's highest diversity forests.

Plants cope with the environment by displaying large phenotypic variation. Two spectra of global plant form and function have been identified: a size spectrum from small to tall species with increasing stem tissue density, leaf size, and seed mass; a leaf economics spectrum reflecting slow to fast returns on investments in leaf nutrients and carbon. When species assemble to communities it is assumed that these spectra are filtered by the environment to produce community level functional composition. It is unknown what are the main drivers for community functional composition in a large area such as Amazonia. We use 13 functional traits, including wood density, seed mass, leaf characteristics, breeding system, nectar production, fruit type, and root characteristics of 812 tree genera (5211 species), and find that they describe two main axes found at the global scale. At community level, the first axis captures not only the 'fast-slow spectrum', but also most size-related traits. Climate and disturbance explain a minor part of this variance compared to soil fertility. Forests on poor soils differ largely in terms of trait values from those on rich soils. Trait composition and soil fertility exert a strong influence on forest functioning: biomass and relative biomass production.

Climate change and increasing availability of resources such as carbon dioxide are modifying forest functioning worldwide, but the effects of these changes on forest structure are unclear. As additional resources become available, for example, through CO2 fertilization or nitrogen deposition, large trees, with greater access to light, may be expected to gain further advantages. Conversely, smaller light-suppressed trees might benefit more if their light compensation point changes, while bigger trees may be the most negatively impacted by increasing heat and drought. We assessed recent changes in the structure of Earth's largest tropical forest by analysing 30 years of Amazonian tree records across 188 mature forest plots. We find that, at a stand level, trees have become larger over time, with mean tree basal area increasing by 3.3% per decade (95% CI 2.4; 4.1). Larger trees have increased in both number and size, yet we observed similar rates of relative size gain in large and small trees. This evidence is consistent with a resource-driven boost for larger trees but also a reduction in suppression among smaller trees. These results, especially the persistence and consistency of tree size increases across Amazonian forest plots, communities and regions, indicate that any negative impacts of climate change on forests and large trees here have so far been mitigated by the positive effects of increased resources.

Several studies have documented dominance by few species in Amazonian forests. Dominant species tend to be either locally abundant (local dominants) or regionally frequent (widespread dominants) but rarely both (oligarchs). Here, we explore relationships between dominance and functional traits. We ask whether: (i) dominance is associated with specific functional profiles and (ii) dominance patterns (local vs. widespread dominants) are associated with different functional traits. We combined census data from 503 forest inventory plots across four lowland forest habitats in western Amazonia with trait information for ~2600 tree species, encompassing data collected in the focal plots and data from published sources. We considered traits that relate to leaf, wood, seed and whole‐plant strategies: specific leaf area (SLA), leaf area (LA), N content per unit leaf mass (LN), wood density (WD), seed mass (SM) and maximum diameter at breast height (DBH max ). Our results reveal that dominant species display different trait combinations depending on the habitat type. Taller dominant species exhibit higher regional frequency, associated with higher dispersal ability and lower local abundance, likely due to negative density dependence. Greater SM contributes to higher regional frequency of dominant species via greater dispersal by birds and mammals and seedling survival. Finally, traits related to resource conservation strategies, such as lower SLA, LA, LN and greater WD, favour higher local densities across most habitats, while the opposite pattern was linked to higher regional frequency. Synthesis . Our findings reveal that (i) dominance is associated with different functional traits depending on the habitat type, and (ii) different functional trait values define distinct dominance patterns. Our study exemplifies the potential of trait‐based approaches to illuminate the ecological mechanisms that may underlie dominance in tropical forests. Finally, accounting for both local abundance and regional frequency when studying dominance is likely to improve our understanding and forecasting of how different species will respond to global change drivers in western Amazonia.

Recent evidence suggests that the ecological footprints of pre-Columbian Indigenous peoples in Amazonia persist in modern forests. Ecological impacts resulting from European colonization c. 1550 CE and the Amazonian Rubber Boom c. 1850 to 1920 CE are largely unexplored but could be important additive influences on forest structure and tree species composition. Using environmental niche models, we show the highest probabilities of pre-Columbian and colonial occupation sites, and hence human-induced ecological influences, occurred in forests along rivers. In many areas, the predicted pre-Columbian and colonial distributions overlap spatially with the potential for superimposed ecological influences. Environmental gradients are known to structure Amazonian vegetation composition, but they are also strong predictors of past human influence, both spatially and temporally. Our comparisons of model outputs with relative abundances of Amazonian tree species suggest that pre-Columbian and colonial-period ecological legacies are associated with modern forest composition.

Despite the progress in the measurement and accessibility of plant trait information, acquiring sufficiently complete data from enough species to answer broad-scale questions in plant functional ecology and biogeography remains challenging. A common way to overcome this challenge is by imputation, or 'gap-filling' of trait values. This has proven appropriate when focusing on the overall patterns emerging from the database being imputed. However, some applications force the imputation procedure out of its original scope, using imputed values independently from the imputation context, and specific trait values for a given species are used as input for computing new variables. We tested the performance of three widely used imputation methods (Bayesian hierarchical probabilistic matrix factorization, multiple imputation by chained equations with predictive mean matching, and Rphylopars) on a database of tropical tree and shrub traits. By applying a leave-one-out procedure, we assessed the accuracy and precision of the imputed values and found that out-of-context use of imputed values may bias the estimation of different variables. We also found that low redundancy (i.e. low predictability of a new value on the basis of existing values) in the dataset, not uncommon for empirical datasets, is likely the main cause of low accuracy and precision in the imputed values. We therefore suggest the use of a leave-one-out procedure to test the quality of the imputed values before any out-of-context application of the imputed values, and make practical recommendations to avoid the misuse of imputation procedures. Furthermore, we recommend not publishing gap-filled datasets, publishing instead only the empirical data, together with the imputation method applied and the corresponding script to reproduce the imputation. This will help avoid the spread of imputed data, whose accuracy, precision, and source are difficult to assess and track, into the public domain.

Interactions between host plants, insect herbivores, and parasitoids are fundamental to understanding ecosystem dynamics. In the present study, we describe the network structure of assemblages composed of host plants, galling insects, and micro-hymenopterans in four vegetation types in two protected areas in the Peruvian Amazon. In total, we recorded 13 host plant species, 20 species of galling insects and 31 taxa of micro-hymenopterans. The network of host plants and galling insects in the white sand-wet forest was very specialized, characterized by low connectance, low linkage density and higher modularity than expected based on the null model, compared to other vegetation types. The galling-micro-hymenopteran interactions in the terra firme and white-sand wet forest were characterized by lower connectance, lower linkage density and higher modularity than expected based on the null model. Our findings show that the high specialization in the plant-gall insect and gall insect-micro-hymenopteran networks reported in other biomes is also found in the Peruvian Amazon.

Understanding the capacity of forests to adapt to climate change is of pivotal importance for conservation science, yet this is still widely unknown. This knowledge gap is particularly acute in high-biodiversity tropical forests. Here, we examined how tropical forests of the Americas have shifted community trait composition in recent decades as a response to changes in climate. Based on historical trait-climate relationships, we found that, overall, the studied functional traits show shifts of less than 8% of what would be expected given the observed changes in climate. However, the recruit assemblage shows shifts of 21% relative to climate change expectation. The most diverse forests on Earth are changing in functional trait composition but at a rate that is fundamentally insufficient to track climate change.

Species' traits and environmental conditions determine the abundance of tree species across the globe. The extent to which traits of dominant and rare tree species differ remains untested across a broad environmental range, limiting our understanding of how species traits and the environment shape forest functional composition. We use a global dataset of tree composition of >22,000 forest plots and 11 traits of 1663 tree species to ask how locally dominant and rare species differ in their trait values, and how these differences are driven by climatic gradients in temperature and water availability in forest biomes across the globe. We find three consistent trait differences between locally dominant and rare species across all biomes; dominant species are taller, have softer wood and higher loading on the multivariate stem strategy axis (related to narrow tracheids and thick bark). The difference between traits of dominant and rare species is more strongly driven by temperature compared to water availability, as temperature might affect a larger number of traits. Therefore, climate change driven global temperature rise may have a strong effect on trait differences between dominant and rare tree species and may lead to changes in species abundances and therefore strong community reassembly.

Unlike most rivers globally, nearly all lowland Amazonian rivers have unregulated flow, supporting seasonally flooded floodplain forests. Floodplain forests harbor a unique tree species assemblage adapted to flooding and specialized fauna, including fruit-eating fish that migrate seasonally into floodplains, favoring expansive floodplain areas. Frugivorous fish are forest-dependent fauna critical to forest regeneration via seed dispersal and support commercial and artisanal fisheries. We implemented linear mixed effects models to investigate drivers of species richness among specialized frugivorous fishes across the ~6,000,000 km 2 Amazon Basin, analyzing 29 species from 9 families (10,058 occurrences). Floodplain predictors per subbasin included floodplain forest extent, tree species richness (309,540 occurrences for 2,506 species), water biogeochemistry, flood duration, and elevation, with river order controlling for longitudinal positioning along the river network. We observed heterogeneous patterns of frugivorous fish species richness, which were positively correlated with floodplain forest extent, tree species richness, and flood duration. The natural hydrological regime facilitates fish access to flooded forests and controls fruit production. Thus, the ability of Amazonian floodplain ecosystems to support frugivorous fish assemblages hinges on extensive and diverse seasonally flooded forests. Given the low functional redundancy in fish seed dispersal networks, diverse frugivorous fish assemblages disperse and maintain diverse forests; vice versa, diverse forests maintain more fish species, underscoring the critically important taxonomic interdependencies that embody Amazonian ecosystems. Effective management strategies must acknowledge that access to diverse and hydrologically functional floodplain forests is essential to ensure the long-term survival of frugivorous fish and, in turn, the long-term sustainability of floodplain forests.

1. Leaf and wood functional traits of trees are related to growth, reproduction, and survival, but the degree of phylogenetic conservatism in these relationships is largely unknown. In this study, we describe the variability of strategies involving leaf, wood and demographic characteristics for tree genera distributed across the Amazon Region, and quantify phylogenetic signal for the characteristics and their relationships. 2. Leaf and wood traits are aligned with demographic variables along two main axes of variation. The first axis represents the coordination of leaf traits describing resource uptake and use, wood density, seed mass, and survival. The second axis represents the coordination between size and growth. Both axes show strong phylogenetic signal, suggesting a constrained evolution influenced by ancestral values, yet the second axis also has an additional, substantial portion of its variation that is driven by functional correlations unrelated to phylogeny, suggesting simultaneously higher evolutionary lability and coordination. 3. Synthesis. Our results suggest that life history strategies of tropical trees are generally phylogenetically conserved, but that tree lineages may have some capability of responding to environmental changes by modulating their growth and size. Overall, we provide the largest-scale synopsis of functional characteristics of Amazonian trees, showing substantial nuance in the evolutionary patterns of individual characteristics and their relationships.

Unraveling the mechanisms underlying the maintenance of species diversity is a central pursuit in ecology. It has been hypothesized that ectomycorrhizal (EcM) in contrast to arbuscular mycorrhizal fungi can reduce tree species diversity in local communities, which remains to be tested at the global scale. To address this gap, we analyzed global forest inventory data and revealed that the relationship between tree species richness and EcM tree proportion varied along environmental gradients. Specifically, the relationship is more negative at low latitudes and in moist conditions but is unimodal at high latitudes and in arid conditions. The negative association of EcM tree proportion on species diversity at low latitudes and in humid conditions is likely due to more negative plant-soil microbial interactions in these regions. These findings extend our knowledge on the mechanisms shaping global patterns in plant species diversity from a belowground view.

Amazonia's floodplain system is the largest and most biodiverse on Earth. Although forests are crucial to the ecological integrity of floodplains, our understanding of their species composition and how this may differ from surrounding forest types is still far too limited, particularly as changing inundation regimes begin to reshape floodplain tree communities and the critical ecosystem functions they underpin. Here we address this gap by taking a spatially explicit look at Amazonia-wide patterns of tree-species turnover and ecological specialization of the region's floodplain forests. We show that the majority of Amazonian tree species can inhabit floodplains, and about a sixth of Amazonian tree diversity is ecologically specialized on floodplains. The degree of specialization in floodplain communities is driven by regional flood patterns, with the most compositionally differentiated floodplain forests located centrally within the fluvial network and contingent on the most extraordinary flood magnitudes regionally. Our results provide a spatially explicit view of ecological specialization of floodplain forest communities and expose the need for whole-basin hydrological integrity to protect the Amazon's tree diversity and its function.

AimEcological and anthropogenic factors shift the abundances of dominant and rare tree species within local forest communities, thus affecting species composition and ecosystem functioning. To inform forest and conservation management it is important to understand the drivers of dominance and rarity in local tree communities. We answer the following research questions: (1) What are the patterns of dominance and rarity in tree communities? (2) Which ecological and anthropogenic factors predict these patterns? And (3) what is the extinction risk of locally dominant and rare tree species?LocationGlobal.Time period1990-2017.Major taxa studiedTrees.MethodsWe used 1.2 million forest plots and quantified local tree dominance as the relative plot basal area of the single most dominant species and local rarity as the percentage of species that contribute together to the least 10% of plot basal area. We mapped global community dominance and rarity using machine learning models and evaluated the ecological and anthropogenic predictors with linear models. Extinction risk, for example threatened status, of geographically widespread dominant and rare species was evaluated.ResultsCommunity dominance and rarity show contrasting latitudinal trends, with boreal forests having high levels of dominance and tropical forests having high levels of rarity. Increasing annual precipitation reduces community dominance, probably because precipitation is related to an increase in tree density and richness. Additionally, stand age is positively related to community dominance, due to stem diameter increase of the most dominant species. Surprisingly, we find that locally dominant and rare species, which are geographically widespread in our data, have an equally high rate of elevated extinction due to declining populations through large-scale land degradation.Main conclusionsBy linking patterns and predictors of community dominance and rarity to extinction risk, our results suggest that also widespread species should be considered in large-scale management and conservation practices.